Advances in Marine Biology, Volume 18

Advances in

MARINE BIOLOGY VOLUME 18

This Page Intentionally Left Blank

Advances in

MARINE BIOLOGY VOLUME 18 Edited by

J. H. S. BLAXTER Dunstaffnage Marine Research Laboratory, Oban, Scotland

SIR FREDERICK S. RUSSELL Ptymouth, England

and

SIR MAURICE YONGE Edinburgh, Scotland

Academic Press

1980

A Subsidiary of Harcourt Brace Jovanovieh, Publishem

London

New York

Toronto

Sydney

San Francisco

ACADEMIC PRESS

24-28

mc.

(LONDON) LTD.

OVAL ROAD

LONDON N W 1 7DX

U.S. Edition published by ACADEMIC PRESS INC.

111 FIFTH

AVENUE

NEW YORK, NEW YORK

10003

Copyright @ 1980 by Academic Press Inc. (London) Ltd.

All rights reserved

b 1 ) PART O F THIS BOOK MAY B E REPRODUCED I N ANY FORM BY PHOTOSTAT, 1\1 ICROFILM, OR ANY OTHER MEANS, WITHOUT WRITTEN PERMISSION FROM

T H E PUBLISHERS

British Library Cataloguing in Publication Data Advances in marine biology. Vol. 18 1. Marine biology I. Blaxter, J. H. S. 11. Russell, Sir Frederick Stratten 111. Yonge, Sir Maurice 574.92

QH91

63-14040

ISBN 0-12-026118-9

Printed in Great Britain by John Wright & Sons Ltd, The Stonebridge Press, Bristol

THE BIOLOGY OF MYSIDS AND EUPHAUSIIDS

JOHNMAUCHLINE The Dunstaffnage Marine Research Laboratory P.O. Box No. 3, Oban, Argyll, Scotland

Dedicated to my wife Isobel and to John, Rowena and Jenqer

PREFACE A N D ACKNOWLEDGEMENTS Euphausiids and mysids are linked together historically because of mperficial similarities-they were originally known as the Schizopoda. The euphausiids, however, are now recognized as decapod and the mysids as peracarid crustaceans. This early linkage of the two Orders has resulted in a continuing interest in both of them by modern workers. This volume should encourage the dual interest which has benefited earlier work and will be of considerable advantage in the future. Many colleagues throughout the world have encouraged and helped in the production of this work. I wish to acknowledge the helpful letters from and/or discussions with Dr M. S. de Almeida Prado, Dr M. Bacescu, Mr A. de C. Baker, Dr E. Brinton, Dr I. Everson, Dr R. R. Makarov, Dr M. Murano, Dr T. Nemoto and Dr J. R. Sargent. It is not possible to mention everybody but I would like to thank them all most sincerely. Mr R. I. Currie and my colleagues within the Dunstaffnage Marine Research Laboratory have in many cases been patient listeners and I thank them for their advice and criticism. Our Librarians, Miss E. Walton and Miss R. A. Gow, have enthusiastically sought papers from obscure sources and I am very grateful indeed to them for their perseverance. I owe much to Miss A. E. Leitch, Mrs E. MacDougall and Miss C. A. Keenan who have done much of the typing of the manuscripts. My greatest debt of gratitude is to my assistant Mrs Catherine M. Webster who has been involved in most aspects of this work, both practical and secretarial. Finally, it is once again a pleasure to acknowledge the helpful comments and criticisms of the Editors, Sir Frederick S. Russell, Sir Maurice Yonge and Dr J. H. S. Blaxter.

JOHN MAUCHLINE

vii

This Page Intentionally Left Blank

CONTENTS PREFACE AND ACKNOWLEDGEMENTS . .

. .

..

. .

vii

..

. .

3

2. The species of mysids and key to genera. . . .. 3. The larvae and reproduction

.. . . ..

39

. .

66

. . .. . .

81 117

8. Physiology and responses to physical-chemical para.. .. . . . . meters of the environment

140

Part 1

The Biology of Mysids 1. Introduction.

. .

. .

. .

4. Vertical distribution and migration

. .

.. . . . .

. . .. ..

..

. .

.. ..

. .

5. The gut, food and feeding . 6. Chemical composition

. .

7. Internal anatomy. .

. .

..

. . ..

. . . .

..

11. Geographical distribution .

. .

..

. .

.. . .

9. Behaviour

. .

10. Population dynamics 12. Predators and parasites

13. Mysids in the marine economy

References .

.

..

. .

.. ..

..

98

.. . . . .

. . 187

..

. . 236

..

..

..

..

..

. . 264 . . 320

..

Appendix I. Taxonomic list of lysidacea

.

.

..

6

. .

I 60

220 258

Appendix 11. Classified list of literature for each genus

343

Appendix 111. Classified list of literature for geographical . . .. . . . . . . regions . . . .

364

ix

Part I t T h e Biology of Euphausiids

. . . .

. .

. .

. .

. .

. .

. .

3. Distribution and synonymy

.. . .

. .

. .

. .

. .

. .

. .

. . . .

. . . .

. . . . ..

. . . . .

. . . . .

. .

1.

Introduction.

2.

The species of krill .

4. The larvae. .

. .

. .

. . . . . .

373

420

. .

. . . . . . .

. .

. . 515

. .

. . 527

. . . .

.. . .

542

..

. .

..

596

.

. .

. .

..

624

. . . .

. . 639 . . 663

. .

. . . . . .

.

679

. .

. .

. .

681

5. Vertical distribution and migration . . . . 6. Food and feeding . .

. .

..

8. Vision and bioluminescence

. .

7.

9.

Chemical composition

Internal anatomy and physiology

10. Growth, maturity and mortality .

.. 12. Predators and parasites . . 11.

Ecology of distribution

.. . .

13. Euphausiids in the marine economy

References. .

. .

. .

. .

Addendum to Biology of Euphausiids Addendum t o Biology of Mysids Taxonomic Index. .

..

. .

. .

. .

. .

Cumulative Index of Titles

. . . .

Subject Index

Cumulative Index of Authors

X

..

.

. . . . . . .

375 384 430 443 459 478 484 490

554

PART ONE

THE BIOLOGY OF MYSIDS

This Page Intentionally Left Blank

CHAPTER 1

INTRODUCTION Species of the Order Mysidacea are shrimp-like animals that occur in vast numbers in coastal regions of the world. They are often overlooked during sampling programmes because they are not sampled efficiently by either conventional benthic or pelagic samplers. A great many species inhabit water of a few centimetres to a few metres in depth, in the environment that lies between the tidal and strictly sublittoral. Many species swim in the water layer immediately above the sediment, others rest on the surface of the sediment, and a few species actually burrow into the sediment. I n consequence, a variety of sampling methods are required, the most important of which is the towing of plankton nets mounted in frames which will slide over the surface of the sediment. This is an equally valuable method for catching many offshore species living on the continental shelves of the world at depths as great as 200 m. I n addition to the species associated with the sea bed, there are many strictly pelagic species. Some of these live in coastal regions and others are oceanic, a few living in the meso- and bathypelagic environments. The meso- and bathypelagic species are well known, especially species of the genus Gnathophausia, some of which attain a large body size and are easily seen in samples of deep plankton. The shallower living epipelagic species are all smaller, less than about 30 mm in body length, are relatively scarce in samples and not so easily noticed among the mass of euphausiids, decapods, copepods and other organisms that occur in such samples. Mysids are a highly adaptive group. They inhabit all regions of the oceans to depths as great as 7210 m, where Belyaev (1966) records the capture of Amblyops magna in the Kuril-Kamchatka Trench. There are many brackish water species and a few species that occur in freshwater environments. Further species have become adapted to the specialized environments of caves and wells, while yet others live in commensal association with animals such as sea anemones. Superficial similarities exist between the mysids and the euphausiids and resulted in the Mysidacea and Euphausiacea being grouped

4

THE BIOLOUY OF MYSIDS

together in the early literature as the Schizopoda (Mauchline and Fisher, 1969). The euphausiids are now recognized as Decapoda. The Mysidacea are placed in the Class Eumalacostraca, Subclass Peracarida which contains the Orders Cumacea, Tanaidacea, Isopoda, Amphipoda, Mysidacea and a small Order Spelaeogriphacea created by Gordon (1957, 1958, 1960) to contain some peculiar subterranean shrimps. A full historical account of the problems of classification of mysids and euphausiids is given in Mauchline and Fisher (1969) and discussed in a broader context by Raw (1955). The earliest published record of a mysid is in 1808 and Tattersall and Tattersall (1951) provide an excellent historical account of the nineteenth century literature describing these animals. Their monograph emphasizes British species although it does refer to certain aspects of other species, especially bionomics. Gordan (1957) produced a bibliography of the Order listing over 1100 references up to the end of 1955. Gordan’s bibliography is divided into three parts. The first contains an alphabetically arranged list of publications. The second part lists the genera and species of mysids and details the relevant literature. The third part contains classified lists of references under the following subjects : general references ; taxonomy, morphology, embryology, life history and growth ;physiology ; behaviour ; symbiosis, including commensalism and parasitism. She refers, in the last sections of this part, to literature pertinent to the following geographical regions : Freshwater and brackish water distribution : Europe, America, Asia, Australia and New Zealand, Japan, Africa and the Arctic. Oceanic distribution : Atlantic Ocean, Mediterranean, Indian Ocean, Pacific Ocean, Arctic Ocean and Antarctic Ocean. Coastal distribution : Europe (Atlantic coasts), the Mediterranean (including the Adriatic and Black Seas), east coast of North America, Caribbean Sea (including Gulf of Mexico and Bermuda), east coast of South America, the coasts of Africa (including Arabia, Iran, India and Burma), Australasia (including Malay Peninsula, Indo-China, Indonesia, the Philippines Australia and New Zealand), east coast of Asia (including China, Formosa, Korea, Japan and the Soviet Union), Oceania (including Micronesia, Polynesia, Fiji Islands and the Hawaiian Islands), west coast of North America, west coast of Central America, west coast of South America, Arctic Ocean (including Greenland, Iceland, Spitzbergen and northern coasts of North America, Europe and Asia) and the Antarctic Ocean. This detailed classification of the literature has made Gordan’s bibliography extremely valuable. The late Dr W. D. Clarke, through

1.

INTRODUCTION

5

correspondence with specialists in the group, identified omissions from and additions to the original bibliography and produced supplementary lists of new references up to 1969 (Holthuis, 1967). Over 700 such references are combined with Gordan’s original reference list and published by Beeton and Clarke (1973), an alphabetical list now consisting of 1852 publications. They do not reproduce Gordan’s subject indices and give no ancillary information on the subject content of the 723 additional references. More than 300 of these are used in the present work and are entered in the classified bibliographies in Appendices I1 and 111. Of the remaining 400 references, some 150 are either general textbooks, fauna lists etc. or are quoted within other references used here. Another 75 publications, issued before 1900, are also in this category. Some 70 papers refer to studies on freshwater environments and most have been superseded by literature quoted here. About 30 references are to theses and internal reports, data from which have usually been published in a more formal way elsewhere. This leaves about 75 papers of the original 723, published predominantly between 1900 and 1950, that have not been obtained during the course of the present work; the titles of the great majority of them suggest that they are concerned with the distribution of individual species rather than with other aspects of their biology. The distributions of the majority of species of mysids are not well known and the production of distributional maps for most of the species is considered to be of little value at the present time. Mauchline and Murano (1977) list broad geographical regions of occurrence of all species and a list of pertinent literature for these regions is given in Appendix 111. Supplementary information is given in Chapter 10. This work does not contain a treatise on the taxonomy of mysids and consequently the following publications are required by specialists as companions to this volume: Tattersall and Tattersall (1951), for British and North Atlantic species ; Tattersall (1951), for American species ; Ii (1964a) and the papers of Murano, for Japanese and western Pacific species; Pillai and some papers of Nouvel, for Indian Ocean species; the papers of Bacescu, Brattegard and 0. S. Tattersall, for species in regions specified in the titles of the papers. Access to other relevant literature is easily obtained through the references quoted by Mauchline and Murano (1977).

CHAPTER 2

THE SPECIES OF MYSIDS AND KEY TO GENERA There are 780 known species of mysids distributed between about 120 genera. Each species is listed by Mauchline and Murano (1977) in the World List with brief notes on their occurrence and a reference to a good description in the literature. There are several corrections and additions to this list and these are set out in Table 11. Mauchline and Murano do not list synonymy of the various species. Informatioh on this aspect is available in Gordan (1957), the taxonomic papers listed at the end of Chapter 1 and the references quoted by Mauchline and Murano. An abbreviated classification and world list of Mysidacea is given in Appendix I. Individual species are identified through differences in their external morphology, especially the structure of such appendages as the thoracic legs, antenna1 scales, telson and uropods. A key to the genera of mysids is g-iven a t the end of this chapter and amply illustrates the characters used. Identification of many species is difficult because of variation in the characters used and their ontogenetic development. Information on the amount and type of variation encountered within some species is available. Much of this has been published in the early taxonomic and morphological literature quoted by Gordan (1957). The Order is sub-divided into the sub-orders Lophogastrida and Mysida. Many species in the Lophogastrida live in the deep oceanic environment and are large in body size relative to species in the Mysida. The largest mysid known, an individual of Gnathophausia ingens of 351 mm total body length (Clarke, 1961a), belongs to this sub-order. Species of the Lophogastrida, with the exception of those in the genera Paralophogaster, have adult body lengths ranging from 17 to 350 mm; species of Paralophogaster are smaller, having a range of body length of 6-20 mm. The majority of species of the Mysida have adult body lengths of less than 15 mm (Mauchline, 1972). Many morphological characters of lophogastrid mysids vary (Fage, 1940, 1941, 1942, 1952; Banner, 1954b), and to such an extent that further investigations of Eucopia species may result in a’ revision of the genus. The variation of some of the taxonomic characters in a

2. THE

SPECIES O F MYSIDS

7

few species of Mysida has been examined. The form of the antennal scale and its relative proportions is frequently an important character used to distinguish species; variation is present within species as shown by Mauchline (1971~)for the closely related P. JEezuosus and P. neglectus and by Williams et al. (1974) for Neomysis americana. The telson of the species in the genus Mysis has a cleft a t its distal end (Fig. 8.17) and such forms of telson occur in other genera. The depth, width and armature of the cleft not only varies between species but also within species, as demonstrated by Holmquist (1949, 1959b). The number of lateral spines on such telsons increases with increasing body size of the animal as shown in Fig. 1 for the two Praunus species and as described in Gastrosaccus vulgaris by Matsudaira et al. (1952). Similar correlations between numbers of spines on the margins of telsons and of both the endopod and exopod of the uropods on the one hand, and increasing body size on the other, have been found in Anchialina m-dagascariensis and A . sanzoi by Nouvel (1969), in Pseudomma afine by Wigley and Burns (1972), in Tenagomysis chiltoni by Hodge (1964), in Neomysis americana by Williams et al. (1974) and in Xiriella clausii by Genovese (1957). Geographi.ca1 variation in the ranges in numbers of spines on the lateral margins of the telsons of both the Praunus species are evident when the Danish and Scottish populations are compared (Mauchline, 197lc). Such variations in several morphometric relationships in Neomysis americana are demonstrated by Williams et al. (1974). Casanova (1977) showed that the eyes of Eucopia hanseni contain different numbers of ommatidia in Atlantic as compared with Mediterranean individuals. Juveniles of closely related species are often indistinguishable. This is true in the two species of Praunus discussed previously; the antennal scales have the same proportions (Mauchline, 1971c) and the lateral margins of the telsons are armed with the same range in numbers of spines (Fig. 1). The patterns of groupings of spines on the margins of the telson or uropods can be diagnostic of some species but the distinctive groupings are often not present in the juveniles. This is instanced by the study of Genovese (1956) on the outer margins of the exopod of the uropod of Siriella clausii (Fig. 2). Juvenile Lophogaster typicus have pectinate edges to the anterior region of the carapace and ventral margins of the abdominal pleura; this ornamentation is absent in the adult (Sars, 1916; Tattersall and Tattersall, 1951). The changing proportions of the body of Gnathophuusia ingens were examined by Clarke (1961a). The rostrum of juveniles, in common with that of many other crustaceans, is longer relative to body length

c

32 r

28

24 c 0 In

A

-

A

-

In

-

A A A

A AA

-

E 0

J

t

16-

12 -

A

A

:A

A A A

A A A f A

A

t

I

I

A

A

A

4

A

4

AA

A

A A

t

A%

&A

A Aft

t A

2 LJ

A

B 4%

f 4i

S

L

& hh

t

A A

A A :

A

.ftA t .tf *$!A t%*A: % : @A ~&%%:% 4% A

A

:

A A :

t

c"

A

A

: t

* 4Y *

t A

A

t

:

*

20Q

f

A

Q)

C ._

:

A A

c

Q) In

A

A A

it

Att

:

:A

PA::

A

2

AA

2A

.

A

A

A

A

A

A

A

A

2

A

A

A

A

A

A

24

A

A

2b 2

A

A

A

A

A

A

A

%

A

A

A A

t f A

A A

bA fB4 A t

-

B

t

A

S

8-

I 3

r

4:

n

AA

I

:

A t

A

A

I

5

I

7

I 9

I

I 11

I

I 13

I

I 15

I

I 17

I

I 19

1

I 21

I

I 23

I

J 25

FIG.1. Number of lateral spines on the telson of Praunwflexwsus (solid triangles) and P . negleetw (open triangles)related to total body length.

2. THE

SPECIES OF MYSIDS

9

FIG.2. Development of spines on the outer edge of the right exopod of the uropod of Skiella clausii. Body lengths of individuals are from left to right: 9, 8.8 mm; 9,8.4 mm; $2, 8.3mm; 3, 8.1mm; d, 7.9mm; 8 , 7.9mm; 9, 7.4mm; 9, 6.6mm; d, 5.6 mm (after Genovese, 1956).

then in fully mature animals. Various biometrical relationships between measurements of the body, rostrum, eye, appendages, telson and uropods are used by Holmquist (1956)in her study of seven species in the genus Boreomysis, The occurrence of secondary sexual characteristics in the adults provides further characters of taxonomic value, especially the detailed form of modified pleopods of males. These are used in the generic key at the end of this chapter. Chemical methods of distinguishing species and populations of mysids have not been investigated in any detail. Populations of the freshwater Mysis relicta exhibit isoenzyme polymorphism (Fiirst and Nyman, 1969). Homogenates of whole M . relicta were subjected to horizontal zone-electrophoresis on starch gel, stained for esterases, and found to contain two zones of activity, A range of patterns was obtained that suggested that five allelic genes producing 15 possible phenotypic combinations were involved. The populations could be separated on the basis of these electrophoretic patterns. One molecular form of esterase was found to occur in M . mixta and Praunus inermis and two in P. Jlexuosus and Neomysis integer by Nyman and Westin (1969). The different patterns obtained allowed electrophoretic separation of the species. Holmquist (1957a) and Nouvel (1964~)found aberrant individuals of Boreomysis rostrata, Mysis oculata, Praunus Jexuosus, P. neglectus, P. inermis and Neomysis integer. Holmquist suggested that in one

10

THE BIOLOGY O F MYSIDS

locality investigated by her, an industrial chemical effluent might cause abnormalities, although she also considered a general source to be genetical. Nouvel considered such abnormalities to be the result of regeneration after injury. Mysids are capable of regenerating damaged regions as originally demonstrated by Przibram (1901) in Hemimysis Zamornae. Mauchline (unpublished) noticed healed but malformed abdomens in Praunus species in Scotland, as well as the more minor abnormalities of telsons and antenna1 scales described by Holmquist and Nouvel. It seems most likely, as Nouvel suggests, that these are a result of damage inflicted, presumably by predators such as fish and amphipods, and the resultant regeneration. These aberrant individuals are relatively rare in a population and the species to which they belong can usually be identified. Sexual intermediates (intersexes) were first noted by Kinne (1955) in Neomysis integer. Three individuals had feminine characteristics in addition to the fourth pair of pleopods being modified as in the males. Two of these animals had a normally developed marsupium containing, developing embryos. Holmquist (1957a) also recorded intersexes of the same form in this species. I n addition, however, she observed one individual, badly damaged, with the male genital organs a t the bases of the fourth thoracic legs and with a t least one pair of brood lamellae. Another form of variation between populations of the same species is in the overall body size. The maximal or modal body size of a species can vary between different localities within one geographical region, as shown by the data of Williams et al. (1974) on Neomysis americana. Body size also varies within a species when its latitudinal range is considered. Casanova (1977) shows that Eucopia hanseni is of a larger body size in the Bay of Biscay than in the Mediterranean. Mauchline (1970b), however, found two distinct forms of Mysidopsis gibbosa in the Clyde Sea Area, Scotland, which differ from each other in body size, general form and the depth at which they occur. The normal smaller form, with a range of adult body length of P 6 m m , lives in bays usually over sand in depths ranging from 1 to 5 m. The body is as illustrated by Tattersall and Tattersall (1951) with the abdomen sigmoid in shape in lateral view. The larger form, measuring 5-8mm when adult, occurs a t depths of approximately 50m over mud or gravel. Its abdomen is straight and it closely resembles young M . didelphys from which it can be distinguished by the form of the telson. Analogous large-sized races of Holmsiella afinis and H . anomala from the Pacific Ocean are discussed by Murano (1970c, 1976). Again, the large-sized race occurs deeper than the small-sized race. Murano

2. THE

11

SPECIES OF MYSIDS

describes minor morphological differences between the two races of H . afinis. So far, the variations that have been discussed are considered to be of a minor nature and to be consistent with the normal patterns of variation expected within species. There are, however, documented variations within other species that have been judged significant enough to require the establishment of sub-species. A list of such sub-species is given in Table I. There is aome inconsistency between the description of forms or races of a species on the one hand and the establishment of sub-species on the other. For instance, Murano (1970~) states that the large form of Holmsiella afinis has larger eyes relative to its body size than the smaller form and there are also other morphological differences. Two of the sub-species listed in Table I also differ from their respective species in minor morphological characters and the size of their eyes : they are Boreomysis rostrata var. Illig and Mysidetes posthm microphthulma. TABLEI. LIST OF SUB-GENERA AND SUB-SPECIES OF MYSIDS Acanthomysis lonqicornis &bra Bacescu, 1972 Acanthomysis sculpta huda Banner, 1948 Anchialina typica orientalis Nouvel, 1971 Anisomysis (Anisomysis) kunduchiana Bacescu, 1975 A n k m y s i s ijimi estafricana Bacescu, 1973c Anisomysis mixta aiwrtralis Zimmer, 1918; Bacescu, 1973b Anisomysis (Paranisomysis) mark rubri Bacescu, 1973d Antromysis (Anophelina) anophelinae Bacescu and Orghidan, 197 1 Antromysis (Parvimysis) almyra Brattegard, 1969 ; Bowman, 197713 A n t r m y s i s (Parvimysis) bahamensis Brattegard, 1969 ; Bowman, 1977b Antromysis (Surinamysis) americana Bowman, 1977b Boreomysis californica longirostris Birstein and Tchindonova, 1958 Boreomysis rostrata japonica Ii, 1964a Boreomysk rostrata orienta1l;e Ii, 1964a Boreomysis rostrata var. Illig, 0. S. Tattersall, 1955 Boreomysis tridens lobata Nouvel, 1942 Diamysis bahirensis mecnikowi (Czerniavsky, 1882); Bacescu, 1954 Cfastrosaccw lobatw var. armata Nouvel, 1951b Cfastrosaccw sanctw widhalmi (Czerniavsky, 1882); Bacescu, 1970a Hemimysis lamorme mediterranea Bacescu, 1941 Hemimysis lamornae pontica (Czerniavsky, 1882); Bacescu, 1954 Heteromysis bermudensis ceaari Bacescu, 1968b Leptomysis apiops banyubnsie Bacescu, 1966 Leptomysis sardiccl pontica Bacescu, 1938 Mesopodopsis africana mmkgascariensis Nouvel, 1978 Metamysidopsis elongata atlantim Bacescu, 1968c mtd.

12

THE BIOLOGY OF M Y S I D S

TABLEI.-(cont.)

Mysidella t y p b mauritanica Bacescu (in press) Myaidetea poathon microphthalma Rustad, 1930 Mysidopst mortenaeni c u b a n b Bacescu, 1968c M y a t (Auricomyak)atenolepis Holmquist, 1959b Nouvelia natalemis mombaaae Bacescu and Vasilescu, 1973 Paramysia baeri biapinoaa Martinov, 1924; Bacescu, 1954 Paramysir, keaaleri sarsi (Derjavin, 1925) ; Bacescu, 1938, 1954 Paramyah lucwtri8 tamitica (Martinov, 1924) ; Bacescu, 1954 Paramysis lacuatrk turcia Bacescu, 1949; Bacescu, 1954 Siriella jaltensia brooki Norman, 1886; Tattersall and Tattersall, 1951 Sirielh jaltewk craasipes (G. 0. Sars, 1877); Bacescu, 1938 Siriella jaltemk grm'lipes Nouvel, 1942 Sirielh japonica izuensis Ii, 1964a Siriella japonica sagamiemis Ii, 1964a Siriella vulgaris rostrata W . M . Tattersall, 1951 Siriella watasei koreana Ii, 1964a Siriella wataaei macropsis Ii, 19648 Tenagomysis (Nouvelia) tanzaniana Bacescu, 1975

Additions .and corrections to the World List published by Mauchline and Murano (1977) are shown in Table 11. The genus Antromysis has been revised by Bowman (197713) and now includes the four subgenera Antromysis Creaser, 1936, Anophelina Bacescu and Orghidan, 1971, Surinamysis Bowman, 1977 and Parvimysis Brattegard, 1969. The full list of species now in this genus is given in Appendix I. It includes the original three species listed by Mauchline and Murano (1977), three new species shown in Table 11, the two species listed by Mauchline and Murano in the genus Parvimysis ( P . almyra and P . bahumensis), and a species in the genus Diamysis, D. americana. Bowman transfers this last species to the genus Antromysis, creating the new sub-genus Surinamysis to receive it. A re-examination of species in the genus Eucopia is required because of the intraspecific variation that has been found in this genus. Likewise, there are probably outstanding taxonomic problems k the gonus Boreomysis. Bacescu (personal communication) questions whether B. richardi Nouvel, 1942, is synonymous with B. vanhoeflenir, Zimmer, 1914. Murano (personal communication) considers that B. tanakai Ii, 1964 is certainly a juvenile form of B. rostrata Ii, 1964 or of B. rostrata orientalis Ii, 1964. Bacescu (personal communication) points out that Potamomysis assimilis Tattersall, 1908 should not be transferred to the genus Diamysis as shown by Mauchline and Murano (1977) since the fourth pleo-

2. THE SPECIES

OF MYSIDS

13

pods of the male differ from those of males in that genus; nor does this species conform to the genus Potamomysis Czerniavsky, 1882. Bacescu suggests that it should be known as Gangemysis assimilis (Tattersall, 1908), this name being used for it by Tattersall and Tattersall (1951). Hatzakis (1977) describes a new species in the genus Haplostylus Kossman, 1880 (Table 11); Mauchline and Murano (1977) wrongly ) the authority for this genus. Species in the quote Bacescu ( 1 9 7 3 ~ as genera Haplostylus and Gastrosaccus are closely similar but can be distinguished by the form of the endopod of the third pair of pleopods of the male (see key to genera). Hatzakis suggests that, on the basis of this character, G. lobatm Nouvel, 1951, G. magnilobatus Bacescu and Schiecke, 1974 and G. normani G. 0. Sars, 1877 should be transferred to the genus Haplostylus. I n the genus Leptomysis, Bacescu (1938), and in a recent personal communication, disagrees with the re-naming of L. sardica G. 0. Sars, 1877 as L. lingvura, f i s t suggested by Illig (1930) and continued by Tattersall and Tattersall (1951). Mauchline and Murano (1977) list three species in the genus Mesomysis. According to Bacescu (personal communication) M . incerta should be transferred to the genus Paramysis as P. incerta (G. 0. Sars, 1895). The other two species, Mesomysis czerniavsky and M . kowatewski have been synonymized with Paramysis lacustris. The genus Nouvelia Bacescu and Vasilescu, 1973 has been re-examined by Bacescu (1975). It now contains three species, all transferred from other genera (Appendix I). These species were previously named (Mauchline and Murano, 1977) as Dozomysis valdiviae (Illig, 1906), Tenagomysis natalensis 0. S . Tattersall (1952) and T. nigeriensis 0. S . Tattersall, 1957. Bacescu considers that T. similis W. M. Tattersall, 1923 may also belong to the genus Nouvelia. No comprehensive world key to the genera of Mysidacea has been published. There are various keys to species within several genera but with the current rate of description of new species most of the keys become quickly out of date. Ii (1964a) provides generic and species keys t o the Japanese mysid fauna, Tattersall and Tattersall (1951) and Nouvel (1950) provide keys for the north-east Atlantic species, and Brunel (1960) to species of the Canadian Atlantic Shelf. Many authors give keys to species within genera and these are among the references listed in the generic bibliography (Appendix 11).The World List of Mauchline and Murano (1977) serves an important function. The genera are listed alphabetically and each species is shown along with its geographical region of occurrence and a t least one source

TABLE

11. ADDITIONSAND CORRECTIONS TO WORLD LIST O F M A U C H L W E AND

Genera and species Antromysis Creaser, 1936 A . juberthiei Bacescu and Orghidan, 1977 A. peckorum Bowman, 1977 A. reddelli Bowman, 1977

Body length (mm)

Zones of occurrence (see Fig. 62)

22N

2.6-3.2

2

caves

Cuba

2 2

caves caves

Jamaica Mexico

Bacescu and Orghidan (1977) Bowman (1977b) Bowman (1977b)

9 9-1 1 10-11

2 2 2

coastal coastal coastal

Brazil Brazil Brazil

Silva (1970a, 1971a) Silva (1972, 1977) Silva (1971b, 1977)

24

1l c

50-585

Kerguelen I

Ledoyer (1977)

3

50-100

S. Africa

Nouvel(1973d)

3a

30-100

Mediterranean Hatzakis (1977)

23s 5-7s

8s

3 4

Ceratomysis Faxon, 1893 C . ericula Ledoyer, 1977

47s

Eryth,rops G. 0. Sam, 1869 E. bidentata Nouvel, 1973

33s

5.5

Haplostylus Kossmann, 1880 H . bacescui Hatzakis, 1977

38N

7-11

Heteromysis S. I. Smith, 1874 H . cotti

(1977)

Latitude (deg)

18N 17N

Bouwmniella Bacescu, 1968 B . atlantica Silva, 1971 B . inarticulata Silva, 1972 B . recifensis Silva, 1971

MURAN0

Habitat or depth (m)

Notes on occurrence

Sources of description

Transfer this species to the genus Heteromysoides Bacescu, 1968

Idiornysis W. M. Tattersall, 1922 I. japonim Murano, 1978

33N

3.3-3.9

7

1-5

Japan

Murano (1978)

Katerythrops Holt and

Tattersall, 1905 R. t k n g u h t a Panampunnayil, 1977 Mesomysis Czerniavsky, 1882 M . incerta G. 0. Sars, 1895

9N

3.5-4.0

8

'coastal

India

Panampunnayil (1977)

Cancel this genus Transfer this species to the genus .Paramy& Czerniavsky, 1882

Metamysidop&, W. M. Tattersall, 1951 M . mamen& Silva, 1970

24s

-

2

coastal

Brazil

Silva (1970b)

Paramy& Czerniavsky, 1882 P . grimmi (G. 0 . Sars, 1895) P . infEata (G. 0 . Sars, 1907)

-

30-40 40

3b 3b

100 8-36

Caspian Caspian

G. 0. Sars (1895) G. 0. Sars (1907)

Pburerythrops Ii, 1964 P . constricta Panampunnayil, 1977 Pseudomysidetes W. Tattersall, 1936 P . cochinensis Panampunnayil, 1977

Sirielh Dana, 1850. S. castellabatensis Ariani and Spagnuola, 1976 S. melloi Silva, 1974

9N

3.5

8

coastal

India

Panampunnayil (1977)

10N

> 3.5

8

coastal

India

Panampunnayil (1977)

40N

10.5-1 1.5

3a

coastal

Italy

9

2

coastal

Brazil

Ariani and Spagnuola (1976) Silva (1974, 1977)

5N-5S

16

THE BIOLOGY O F MYSIDS

of adequate description. Many coastal species occur in restricted localities so that the site of sampling is frequently of help in identifying the species. The key to the genera, appended to the end of this chapter, should be used in conjunction with the World List. The unknown species can be identified to the generic level from the key and descriptions of species within the genus can be found by reference to the World List, taking into account the additions and corrections listed in Table I1 and discussed above. There are two areas of the Key to Genera that are difficult. The four closely related genera Doxomysis, Afromysis, Iimysis and Tenagomysis within the tribe Leptomysini will probably require revision once males of all species have been described (Bacescu, 1968a). Identification of genera within the tribe Mysini is not easy because the principal character used is the form of the maIe pleopods, especially the fourth pair. These are generically distinct and in practice the females can be identified by comparison with the males because large sexual differences in features such as the antennal scale, telson and uropods are rare. In addition, the distributional patterns of chromatophores on the body of live or freshly preserved mysids are often e

FIQ.3. Families Lepidomysidae and Stygiomysidae.A, Spelaeomysis body form showing B, the antennal scale and C, the telson. D, Stygimysis body form showing E, the antennal scale and F, the telson (after Ingle (1972) and Bowman (1976)). e, eye; s, antennal scale.

2. THE SPECIES

17

OF MYSIDS

specific. Identification of juveniles can sometimes be made in this way when other adult morphological characters are undeveloped. This key is a first attempt a t including all genera but areas of it will no doubt require revision when more information becomes available. There are two families of Mysida not included in the generic key. They are the Lepidomysidae and Stygiomysidae, the species of which inhabit caves and wells. Gordon (1958, 1960) and Nath and Pillai (1971) discuss their systematic positions. Ingle (1972) re-examines the Lepidomysidae and groups the species within the single genus Spelaeomysis. These mysids have the body form illustrated in Fig. 3 ; there is no statocyst in the uropods, the eyes are reduced, but the antennal scale is normally developed although small. The Stygiomysidae is represented by a single genus, Stygiomysis. The carapace does not cover the posterior four thoracic segments (Fig. 3), no statocysts are present in the uropods, and both the eyes and antennal scales are reduced. The marsupium consists of seven pairs of lamellae in the Lepidomysidae but the number of pairs of lamellae present in Stygiomysidae has not been determined with certainty; there are at least four and possibly seven pairs.

OF MYSIDS KEYTO GENERA

1. Branchiae present on at Ieast some of the thoracic legs. Pleo-

pods well developed and unmodified in both sexes. No statocyst in endopod of uropod. Marsupium with seven pairs of lamellae (Fig. 4.2) (Sub-order LOPHOQASTRIDA) .. ..

2

Branchiae absent. Pleopods of female reduced, rudimentary (Fig. 4.1); of male variable. Marsupium usually with less than seven pairs of lamellae. Statocyst usually present in endopod of uropod (Sub-order MYSIDA) .. .. ..

8

2. Pleural plates on abdominal segments distinct (Family LOPHOQASTRIDAE)

..

..

..

..

..

..

3

No pleural plates on abdominal segments. Outer margin of antennal scale naked (Fig. 4.3); telson entire (Figs 4.4 and 4.6) (Family EUCOPIIDAE) .. .. .. .. Eucopia 3. Exopod of uropod with suture (Fig. 4.6)

..

Exopod of uropod without suture (Fig. 4.7)

..

..

4

..

..

5

A1

as

u

I

A1

3 su

P ux

0

5

6

FIG.4. Families Eucopiidae, Lophogastridae and Petalophthahnidae. A l , first abdominal segment; as, antenna1 scale; c, carapace; e, eye; 1, lamellee of marsupium; p, rudimentary pleopod; su, suture; t, telson; u, uropod; un, endopod of uropod; ux, exopod of uropod.

2. T H E

4. Long spear-like rostrum present

..

Very small triangular rostrum present 5. Telson of form shown in Fig. 4.8

Telson entire

..

19

SPECIES OF MYSIDS

..

..

.. ..

.. ..

.. ..

Gnathophawia Paralophogaster

..

Ceratolepis

..

..

6

6. Eyes developed; telson and uropods of form shown in Fig. 4.7 .. .. .. .. . . Lophogmter

..

..

Eyes reduced; telson and uropods as in Fig. 4.9 7. Outer margin of antennal scale with 2-4 spines

..

..

7

Chalarmpidum

Outer margin of antennal scale with 9-12 spines Pseudochalaraspidum 8. Statocyst absent (Family

PETALOPHTHALMIDAE)

Statocyst present (Family

MYSIDAE)

I.

..

..

9

..

..

11

9. Body with very extensive spiny armature (Fig. 4.10)

Body without very extensive spiny armature

Ceratomysis

..

..

10

10. Eyes small but stalked and normally shaped (Fig. 4.12)

Petalophthalmus Eyes plate-like or extremely reduced (Figs 4.11, 4.13) Hansenomysis 11. Exopod of uropod with distinct distal suture (Fig. 5.1)

12

Exopod of uropod with the proximal part of the outer margin naked and maxked distally with one or two spines and a transverse articulation (Fig. 5.2); telaon cleft as in Fig. 5.2 Marsupium consists of seven pairs of lamellae (SubFamily BOREOMYSINAE) .. - . .. . . Boreomysis Exopod of uropod undivided

..

..

..

..

..

14

12. Endopod of uropod with distinct distal suture and a strong

spine present on the ventral 'surface close to the statocyst (Fig. 5.3). Marsupium consists of three pairs of lamellae (Sub-Family RHOPALOPHTHALMINAE) .. Rhopulophthalmus Endopod of uropod undivided (Sub-Family SIRIELLINAE)

..

13

20

TEE BIOLOGY OF MYSIDS

17

no. 6. Sub-FamiliesBoreomysinm,Siriellinae,Rhopalophthalminae, Gestrosaccinae and Mysinm. a, precoxa; b, coxa; c, basis; ca, carpus; cp, fused carpopropodus subdivided into 6 perts; d, preischium; da, dectylus; en, endopod; ex, exopod; fm, flexor muscle; is, ischium; k, knee; me, merus; n, nail; 08, oblique articulation; pr. propodus; pr2, propodus composed of two sub-segments;8, statocyst; BU, Butwe; te, transverse articulation.

13. 'Third pair of thoracic legs normal and similar to the more

posterior legs

..

..

..

*.

..

* .

Siriella

Third pair of thoracic legs extremely elongated, almost twice as long as more posterior legs . . .. . . Hemisiriella

2.

21

THE SPECIIES OF MYSIDS

14. One to many spines but no setae on the outer margin of the

exopod of the uropod (Figs 5.4, 5.5, 5.6). Marsupium consists of two pairs of lamellae (Sub-Family QASTROSACCINAE)

15

Outer margin of exopod of uropod with setae but no spines

19

16. Outer margin of exopod of uropod with one or two spines

16

Outer margin of exopod of uropod with more than ten spines

17

16. Outer margin of exopod of uropod naked with one terminal spine (Fig. 6.5) . . .. . . Pseudanchialina

..

,.

Proximal half of outer margin of exopod of uropod naked, marked distally by two spines (Fig. 5.6) .. Paranchialina 17. In male, exopod of pleopod I11 slightly elongated, endopod with many segments (Fig. 5.7) Anchialina

..

..

..

In male, exopod of pleopod I11 greatly elongated and forming a copulatory organ; endopod unsegmented (Fig. 5.8) Bouvmaniella In male, exopod of pleopod I11 greatly elongated, without copulatory organ, and with the cndopod unsegmented (Fig. 6.9) .. .. .. .. .. . . Haplostylus

..

In male, exopod of pleopod I11 greatly elongated and endopod of four segments (Fig. 5.10) . . .. A rchaeomysis In male, exopod of pleopod I11 greatly elongated, endopod with many segments (Fig. 5.11) .. .. . Qaatrosmus Iidla

.

Pleopod of other form

..

..

18. In females, pleopods I-V biramous

In females, pleopods I-V uniramous

..

..

..

..

..

..

..

A rchaeomysis Bowmaniella .. Anchialina Iiella

..

18

I n females, pleopod I biramous, pleopods 11-V uniramous Qaatrosaccus Haplostylus 19. Labrum normal and symmetrical. Mandible with the cutting

lobe not expanded and its edge with spines. Maxillules

22

THE BIOLOGY OF MYSIDS

normal. First thoracic leg with distal segment of endopod normal and without spines on a free distal margin. Marsupium consists of two or three pairs of lamellae. (Sub-Family

..

MYSINAE)

..

..

..

..

..

..

20

Labrum asymmetrical, produced posteriorly into a large plate with two unequal lobes (Fig. 5.12). Mandibles with cutting lobe expanded and without spines (Fig. 5.13). Maxillules with lobes strongly bent inwards. First thoracic leg with last segment of endopod expanded and with a free distal margin armed with spines (Fig. 5.14). Marsupium consists of three pairs of lamellae (Sub-Family MYSIDELLINAE)

..

..

..

..

..

..

..

..

91

20. Third thoracic leg normal and similar to the more posterior legs, with the propodus or fused carpopropodus (Fig. 5.15)

..

..

..

..

21

Third thoracic leg strongly thickened with the carpus and propodus undivided, thickened and armed with spines (Fig. 5.16) (Tribe Heteromysini) . . .. .. .. ..

90

sub-divided

..

..

..

21. Endopods of third to eighth thoracic legs with undivided

carpus marked off from propodus by an oblique articulation (Fig. 5.17) (transverse in Arachnomysis and some species of Pseudomma). Antennal scale with outer margin non-setose (setose in Nipponerythrops) nearly always with a pronounced external spine (scale sometimes absent or represented by a spine). Pleopods 11-V of male well developed and biramous . . .. .. .. telson entire (Tribe Erythropini)

23

Endopods of third to eighth thoracic legs with carpus and propodus fused and sub-divided (Fig. 5.18); no oblique articulation except in Inusitatomysis .. .. ..

22

22. Pleopods 11-V of male usually well developed and biramous

but are rudimentary in Mysidetes. Antennal scale setose all .. .. around, telson variable (Tribe Leptomysini)

59

At least pleopod I1 of male is rudimentary and uniramous, exopod of pleopod IV elongated and modified except in Inusitatomysis where the endopod is elongated and the exopod rudimentary. Antennal scale and telson very variable (Tribe Mysini) . . .. .. .. .. ..

75

2. THE SPECIES

23

OF MYSIDS

..

.. Antennal scale absent or reduced to a spine . .

23. Antennal scale present

..

..

.. ..

..

..

24. Antennal scale with outer margin naked

Antennal scale setose all round 25. Last thoracic segment not elongated

.. ..

.. ..

..

..

24 56 25

Nipponerythrops Thallasomysis

..

..

26

Last thoracic segment greatly elongated making %hecarapace appear very short (Fig. 6.1) .. .. . . Longithorax 26. Visual elements of eye divided into two quite distinct regions .. .. .. .. .. .. . . (Fig. 6.2)

27

Visual elements of eye, when present, not divided into distinct regions . . .. .. .. .. .. ..

28

27. Antennal scale longer, or a t least only slightly shorter, than

antennular peduncle; proximal region of outer margin of scale naked and marked distally by a spine (Fig. 6.3) Euchaetomera Antennal scale about equal in length to the antennular ped.. . . Euchaetomeropsis uncle, setose all round (Fig. 6.4) Antennal scale about half the length of the antennular peduncle and armed with long plumose setae in the distal region only (Fig. 6.5); telson usually of form shown in Fig. 6.6 .. .. .. ., Caesaromysis 8

..

..

Antennal scale about half the length of the antennular peduncle and armed with plumose setae round the distal half (Fig. 6.7). Telson of form shown in Fig. 6.8 . . . . Echinomysis Antennal scale of peculiar form shown in Fig. 6.9

Echinomysides

28. Eyes well developed with normally functioning visual

..

29

Eyes rudimentary with visual elements reduced or absent

46

elements

..

..

..

..

..

..

..

29. Eyes reniform or markedly flattened dorso-ventrally

Eyes more or less globular

.,

..

..

..

.. ..

30 31

FIG. 6. Sub-Family Mysidae, tribe Erythropini. ep, eye-plate; pp, papillifom process ; sp, spinous process.

2.

25

THE SPECIES OF MYSIDS

30. Telson shorter than broad, its apex armed with four strong spines, the lateral margins unarmed (Fig. 6.10) or their . . Erythrops (in part) distal regions serrulated (Fig. 6.11) . .

Telson longer than broad, its apex truncate, armed with 6 spines, the innermost pair being minute, the outer pairs strong; the lateral margins entirely or partially armed with spines (Fig. 6.12) . . .. .. .. HypererytRrops 31. Lateral margins of telson unarmed

Lateral margins of telson armed

.. ..

..

..

..

32

..

..

..

38

32. Apical region of telson p m e d with two or more pairs of

spines with or without plumose setae

..

..

..

33

Apical region of telson armed with one pair of spines (Fig. 6.13) ; antenna1 scale setose without a spine marking distal Heteroerythrops end of naked outer margin (Figs 6.7, 6.14) . . 33. Apical region of telson armed with two pairs of spines as in Fig. 6.15 . . .. .. .. . . Amuthimysis

..

..

Apical region of telson armed with two pairs of spines of .. .. .. .. .. .. other proportions

34

Apical region of telson armed with three pairs of spines Parerythrops (in part) Apical region of telson armed with four pairs of spines

..

37

34. Eyes sub-globular, oval; telson of form shown in Figs 6.10, 6.11 .. .. .. . . Erythrops (in part)

..

..

..

..

..

35

35. No constriction between thorax and abdomen

..

..

3E

Eyes globular

..

..

..

..

Constriction present between thorax and abdomen (Fig. 6.16) .. .. .. .. .. .. Pleurerythrops 36. Pleopod I of male rudimentary

..

Parerythrops (in part)

Pleopod I of male with normally developed exopod and an .. .. .. . . Meterythrops unsegmented endopod Kuterythrops

26

THE BIOLOGY OF MYSIDS

37. Eyes small and imperfectly developed ; antennal scale shorter than antennular peduncle (Fig. 6.17) Teraterythrops

Eyes of normal size; antennal male much larger than antennular peduncle . . .. .. .. Synerythrops (in part) 38. Only apical region of telson armed, with more than four pairs of spines (Figs 6.18, 6.19) .. .. ..

..

Distal half of telson armed with spines 39. Telson of form shown in Fig. 6.19

Telson of form shown in Fig. 6.18

.. ..

..

..

..

39 41

Metamblyops (in part)

..

..

..

40

40. Antennal scale longer than antennular peduncle

Synerythrops (in part) Gibberythrops

Antennal scale shorter than antennular peduncle 41. Telson elongated, triangular (Figs 6.20-22)

Telson linguiform (Fig. 6.23)

..

..

.. ..

..

43. Antennal scale of form shown in Fig. 6.24

Antennal scale of form shown in Fig. 6.25

.. ..

42

Australerythrops

42. Apical region of telson with two long spines (Fig. 6.22)

Apical region of telson without such long spines

..

..

..

43

..

44

Pseuderythrops

..

Pteromysis

44. Endopod of pleopod IV of male elongated and terminating

..

..

..

Endopod of pleopod IV of male not elongated

.. ..

45. Pair of apical plumose setae present on telson

..

in a stout seta

No apical plumose setae on telson

..

..

..

Holmsiella

..

Eoarythrops A tlanterythrops

Metamblyops (in part)

46. Eyes fused to form a single plate or as two immovable plates in contact with each other at their inner margins (Fig. 6.26)

Eyes separste and distinct

..

..

45

..

..

..

47

50

2.

27

THE SPECIES OF MYSIDS

47. Eye plates without a long sharp spinous process on the outer distal corners (Fig. 6.27) .. .. .. . .

Eye plates with lateral spinous process (Fig. 6.26) 48. Pleopo’ds of male biramous

Pleopods of male uniramous

. .. ,

.. ..

.. ..

48

. . Scolamblyops .. ..

..

49

Michthyops

49. Endopod of pleopod IV of male not longer than exopod Pseudomma

Endopod of pleopod I V of male longer than exopod and armed with an extremely long non-plumose seta Parapseudomma 50. Eyes more or less pyriform with definite stalks

Eyes in form of flat plates without definite stalks

.. ..

..

51

..

54

51 Eyes small, imperfectly developed, no visual elements and

a minute papilliform process on the inner lateral surface (Fig. 6.28) .. .. .. .. .. Pseuclumblyops Eyes of similar form (Fig. 6.29), but without the papilliform process . . .. .. .. .. Paramblyops (in part) Eyes larger and of different form

..

..

..

..

52

52. Eyes large, lens-shaped; antennal scale narrow and shorter

than the antennular peduncle ; apical plumose setae present on the telson .. .. .. .. . . Hyperamblyops Eyes smaller, often imperfectly developed ; antennal scale variable in length ; no apical plumose setae on telson Dactylamblyops 53. Eye plates quadrangular, outer corner rounded (Fig. 6.27)

54

Eye plates triangular, outer corner produced into a pointed process (Fig. 6.30) .. .. .. .. .. ..

55

54. Antennal scale long and usually narrow with the unarmed outer margin terminating in a strong spine beyond which . Amblyops the small apex does not extend (Figs 6.31, 6.32)

.

Antennal scale oval with its apex produced well beyond the spine marking the distal end of the unarmed outer . . .. .. .. Amblyopsoides margin (Fig, 6.33)

28

THE BIOLOGY OF MYSIDS

55. Telson as shown in Fig. 6.34

..

..

..

..

Mysimenzies

Telson as shown in Fig, 6.35; antennal scale of form shown in Fig. 6.32 .. .. .. . . Paramblyops (in part) Telson ranging between the triangular forms shown in Figs 6.36 and 6.37; antennal scale variable, in some like Fig. 6.32, in others like Fig. 6.33 . . .. .. Dactylerythrops 56. Visual elements of eye not divided into separate regions

..

57

Visual elements of eye divided into two quite distinct regions (Fig. 6.2) .. .. .. .. Caesaromysis $2 57. Thorax not elongated between first and second thoracic legs

58

Thorax elongated betiween fmt and second thoracic legs leaving a conspicuous limbless region A rachnomysis

..

58. Posterior margins of abdominal segments armed with spines

Chunomysis Posterior margins of .abdominal segments naked 59. Antennal scale setose all round

..

..

Gymnerythrops

..

..

60

Antennal scale with outer margin naked; eyes in the form . . Calyptomma of Iamellae (Fig. 7.1); telson as in Fig. 7.2 . . 60. Pleopods of male well developed, biramous

..

..

..

61

Pleopods of male rudimentary and like those in female. Carpopropodus of endopods of third to eighth thoracic legs . . Mysidetes composed of many sub-segments (Fig. 7.3) . . 61. Telson without cleft

..

..

..

..

Telson cleft, or at least with an apical incision 62. Endopod of uropod armed with spines

..

.. .. 1 .

..

62

..

65

..

63

Endopod of uropod without spines, or with one minute spine Brmilomysis 63. Telson of form shown in Fig. 7.5; endopod of first thoracic .. .. .. . Metamysidopsis leg as in Fig. 7.6

.

Telson of form shown in Fig. 7.7 ; Endopod of first thoracic leg as in Fig. 7.8; eyes large; endopod of uropod without

FIQ. 7. Sub-Family Mysidee, tribe Leptomysini. e eye; ep eye-plate; truberance.

8s

spiny pro-

30

THE BIOLOGY OF MYSIDS

spines except for short rows of 2-7 close to the statocyst .. .. .. .. .. . . Mysideis (Fig. 7.9) Telson of other form

..

..

..

..

..

..

64

64. Endopod of first thoracic leg as in Fig. 7.10 Mysidopsis (in part)

Endopod of first thoracic leg as in Fig. 7.11 65. Eye small or vestigial

..

..

..

..

..

..

..

Leptomysis

..

66

Eye with accessory eye on dorsal lateral side (Fig. 7.12) Dioptromysis Eyes normal in size

..

..

..

..

..

..

67

66. Eyes conical (Fig. 7.13) or reduced with a finger-like process .. . . Pseudomysis (Fig. 7.14); telson of form in Fig. 7.15

Eyes small, sub-quadrangular with or without definite stalks Bathymysis (in part) (Fig. 7.16); telson of form in Fig. 7.17 67. Telson of form in Fig. 7.18

Telson of other form

..

.. ..

..

..

Pseudomysidetes

..

..

..

..

..

.. ..

68. Telson with unarmed apical incision (Fig. 7.19)

Telson cleft

..

..

..

..

..

..

68 69 70

Mysidopsis (in part)

69, Endopod of first thoracic leg as in Fig. 7.10

Endopod of uropod with spiny protuberance on outer edge and unarmed apical incision in telson (Fig. 7.20) Promysis (in part) 70. Cleft of telson not armed with spines

Cleft of telson armed with spines 71. Distal end of telson as in Fig. 7.21

I.

..

..

..

.. ..

Telson of general form shown in Fig. 7.22

..

..

.. ..

71 72

Afromysis (in part) Promysie (in part) Prionomysis

72. Cleft of telson armed with spines and a pair of plumose setae

73

Cleft of telson armed with spines but without plumose setae

74

2.

31

THE SPECIES OF MYSIDS

73. Distal segment of maxilla not expanded but rectangular (Fig. 7.23, 7.31); telson of form in Figs 7.24 and 8.1 Tenagomysis

Distal segment of maxilla expanded, crescent shaped (Fig. 7.25); telson of form in Fig. 7.26 .. . . Afromysis (in part) Distal segment of maxilla triangular (Fig. 7.27); telson as in Figs 7.26 or 7.28 .. .. .. .. . . Doxomysis Distal segment of maxilla as in Fig. 7.29; telson as in Fig. 7.26 .. .. .. .. .. .. .. Iimysis Distal segment of maxilla as in Fig. 7.30; telson as in Fig. 7.26 . . .. .. .. .. .. . . Hyperiimysis Distal segment of maxilla as in Fig. 7.31 ; telson as in Fig. 8.1 Nouvelia Distal segment of maxilla as in Fig. 8.3; telson as in Fig. 8.2 Pseudoxomysis 74. Telson of form in Fig. 8.4

..

..

..

. . Cubanomysis

Telson of form in Fig: 8.5 with long spines on apices Australomysis Telson of form in Fig. 8.6

..

..

Bathymysis (in part)

76. Antennal scale with outer margin naked, armed or unarmed

with spines or dentate

..

..

Antennal scale setose all round

..

..

..

..

..

..

.. ..

76 81

76. Outer margin of antennal scale armed with more than one spine (Fig. 8.7); telson as in Fig. 8.8 Hemimysis (in part)

Outer margin of antennal scale dentate (Fig. 8.9); telson cleft, armed with spines and two plumose setae (Fig. 8.10) Inusitatomysis Proximal region of outer margin of antennal scale naked, marked or unmarked distally by a spine . . .. ..

77

77. Outer margin of antennal scale without distal spine (Fig. 8.11) . .. .. ..

..

78

Outer margin of antennal scale with distal spine (Fig. 8.12)

79

.

..

..

78. Telson triangular, unarmed (Fig. 8.13)

..

..

..

Idiomysis

Telson varying between forms shown in Figs 8.14 and 8.15 Hemimysis (inpart)

FIG.8. Sub-Family Mysidae, tribes Leptomysini and Mysini.

2. THE SPECIES

33

OF MYSIDS

79. Spine on outer margin of antennal scale articulated (Fig. 8.16); telson cleft (Fig. 8.17) . .. .. Praunus

.

..

Spine on outer margin of antennal scale not articulated 80. Telson entire (Fig. 8.18)

..

Telson of forms as in Figs 8.17-8.19

..

..

..

..

I

..

80

..

Katamysis Caspiomysis

..

Paramysis Schistomysis

81. Eyes of complex form or markedly reduced, with or without

pigment

..

..

..

Eyes normally developed

.. ..

.. ..

..

..

..

82

..

..

..

83

82. Eyes very large, divided into a dorsal and ventral region .. Carnegieomysis (Fig. 8.20); telson as in Fig. 8.21

..

Eyes divided as in Fig. 8.22; telson of forms shown in Figs 8.23 and 8.24 . . .. .. .. Anisomysis (in part) Eyes as in Fig. 7.12; cleft of telson armed with two plumose setae and no spines (Fig. 8.25) .. . . Kainommatomysis Eyes reduced in size, ommatidia absent, pigmented or unpigmented; telson triangular with or without two spines on lateral margins (Fig. 8.26) .. Antromysis (in part) Eyes reduced in size, of form shown in Fig. 6.26. Telson of .. .. .. . . Troglomysis form shown in Fig. 10.7

..

84

Telson without cleft or with cleft without plumose setae .. .. .. .. .. .. .. present . .

85

83. Telson with cleft in which two plumose setae are present

84. Telson as in Fig. 8.27; pleopod IV of male of form in Fig. 8.28 .. .. .. .. .. .. . . Arthromysis

Telson as in Fig. 9.1 ; pleopod IV of male of form in Fig. 9.2 Parastilomysis

The remaining genera of the tribe Mysini are separated on the detailed morphology of the pleopods of the males, especially the form of pleopod I V . Pleopods 111 and V are rudimentary, uniramous and like those of the female unless otherwise stated.

i9 11

I9

16

Bra. 9. Sub-Family Mysidae, tribe Mysini.

2.

35

THE SPECIES OF MYSIDS

85. Exopod of pleopod IV of males with: . .. one or 2 segments

.

three, four or five segments six or more segments . .

.. ..

..

..

..

..

..

..

..

..

..

86 88 89

86. Exopod of pleopod IV unsegmented (Fig. 9.13); telson of form in Fig. 9.4; antennal scale as in Fig. 9.5 . Limnomysis

.

Exopod of pleopod IV unsegmented (Fig. 9.6); telson of form in Fig. 9.7; antennal scale as in Fig. 9.8; pleopod V elongated (Fig. 9.9) .. .. .. .. . . Indomysis Exopod of pleopod IV unsegmented (Fig. 9.10); telson and endopod of uropod as in Fig. 9.11 ; antennal scale of form in Fig. 9.12 .. .. .. .. . . Mesacanthomysis Exopod of pleopod IV unsegmented (Fig. 9.13); telson of form in Fig. 9.14; antennal scale of form in Fig. 9.8 ;pleopod I11 biramous, endopod unsegmented but exopod of four segments .. .. .. .. . . Paracanthomysis Exopod of pleopod IV of two segments (Fig. 9.16); telson without cleft, variable in shape but of general form in Fig. 9.17 87. Distal end of antennal scale pointed (Fig. 9.18)

..

87

Neomysis Acanthomysis

Distal end of antennal scale rounded (Fig. 9.19)

88. Exopod of pleopod IV of three segments (Fig. 9.15); telson with shallow cleft or straight edge (Fig. 8.15); antennal scale .. .. .. .. . Diamysis of form in Fig. 9.12

.

Exopod of pleopod IV of three segments (Fig. 9.26); telson with straight edge (Fig. 8.15) ; antennal scale of form in Fig. 9.12 .. .. .. .. .. .. . . Gangemysis Exopod of pleopod IV of three (four) segments (Fig. 9.20); telson varying between forms shown in Figs 9.21 and 9.22; antennal scale of form in Fig. 9.12 . . .. . . Lycomysis Exopod of pleopod IV of three segments (Fig. 9.23); telson of form in Fig. 9.24; antennal scale of form in Fig. 9.12; pleopod I11 biramous with unsegmented endopod and an .. Mesopodopsis exopod of two segments (Fig. 9.25) . . Exopod of pleopod IV of three segments (Fig. 9.27); telson of variable form, without cleft, and usually like Fig. 9.17;

36

THE BIOLOGY OF MYSIDS

antennal scale of general form in Fig. 9.8; pleopod V may be like that of female or elongated . . .. . . Proneomysis Exopod of pleopod IV of three or four segments (Fig. 10.1); telson varying between forms shown in Figs 10.2 and 10.3; antennal scale of form in Fig. 9.12; pleopod I11 uniramous but elongated . . .. .. .. .. . . Mysidium Exopod of pleopod IV with four segments and with large endopod (Fig. 10.4); telson of forms shown in Fig. 10.5; pleopod I11 uniramous and not elongated Antromysis (in part) Exopod of pleopod IV of four segments (Fig. 10.6); telson of form in Fig. 10.7; antennal scale of general form in Fig. 9.8; pleopod I11 (Fig. 10.8) and pleopod V (Fig. 10.9) segmented and elongated . . .. .. .. . . Taphromysis Exopod of pleopod IV of four segments (F'ig. 10.10); telson of form in Fig. 10.11 ; antennal scale as in Fig. 9.12 ; pleopod I11 biramous, with unsegmented endopod and exopod of three segments (Fig. 10.12) . . .. .. . . Nanomysis Exopod of pleopod IV of four segments (Fig. 10.13); telson variable, examples being shown in Figs 10.14-10.18 antennal scale of general form in Fig. 9.12 . . Anisomysis (in part) Exopod of pleopod IV of four or five segments (Fig. 10.19); telson of form in Fig. 9.17; antennal scale of form in Fig. 10.20 ; pleopod I11 biramous, endopod unsegmented and .. .. .. . . Stilomysis exopod of four segments 89. Exopod of pleopod IV of six or seven segments (Fig. 10.21); telson of form in Fig. 8.17; antenna1 scale varying between forms in Figs 9.8 and 10.20; pleopod I11 biramous, endopod

unsegmented and exopod of four to six segments

..

. . Mysis

Exopod of pleopod IV wibh more than seven segments (Fig. 10.22); telson of form in Fig. 10.23; pleopod I11 (Fig. 10.24) and pleopod V (Fig. 10.25) biramous .. Antarctomysis 90. Eyes more or less quadrangular (Fig. 10.26) with the cornea

in the antero-lateral region of the eye-stalk

..

Heteromysoides

Eyes globular or cylindrical, cornea more or less symmetrical relative to the eyestalk. Telson variable (Figs 10.27-10.29) and antennal scale of form shown in Fig. 10.30 . . Heteromysis Neoheteromysis

14

Fw

15

‘P I

19

16

i

I

17 ,

D

8

6

I

w 18

i

22

FIG. 10. Sub-Family Mysidae, tribe Mysini and Heteromysini; Sub-Family Mysidellinae.

38

THE BIOLOGY OF MYSIDS

91. Terminal segment of endopod of first thoracic leg armed with spines (Fig. 10.31); antenna1 scale of form in Fig. 10.30; telson of form in Fig. 10.28 or with a deeper or shallower cleft .. .. .. .. .. .. . . Mysidella

CHAPTER 3

THE LARVAE AND REPRODUCTION Mysids, like other peracaridan crustaceans, carry their embryos in a marsupium within which the entire, although abbreviated, larval development takes place. Young mysids emerge from the marsupium at an early juvenile stage of development. The marsupium is a chamber formed by lamellae or oostegites present on all or some of the second to eighth pairs of thoracic legs. Each of these posterior seven pairs of legs have a pair of lamellae in species belonging to the Sub-order Lophogastrida (Fig. 4.2). I n the Sub-Order Mysida, there are seven pairs of lamellae in species belonging to the Families Petalophthalmidae, Lepidomysidae and also probably Stygiomysidae, although this requires confirmation. The number of pairs of lamellae comprising the marsupium of species belonging to the Family Mysidae of the Sub-Order Mysida vanes; there are seven pairs in the Sub-Family Boreomysinae, three in the Families Siriellinae, Rhopalophthalminae and Mysidellinae, two in the Gastrosaccinae (Fig. 4.1) and two or three in the Mysinae. When only two or three pairs of lamellae are present, they arise from the seventh and eighth and sixth to eighth thoracic legs respectively. The lamellae are normally thin-walled, transparent concave plates fringed with short strong setae that interlock ventrally to form a closed chamber below the thorax. This chamber is normally single but Heteromysis species have two chambers formed side by side because the lamellae fold inwards and upwards in the mid-ventral line so producing lateral chambers (Tattersall and Tattersall, 1951). The eggs emerge within the marsupium from the external genital openings of the oviducts located near the bases of the sixth pair of thoracic legs. The ripe ovary fills the posterior thorax and extends into the abdomen (Fig. 11). Nair (1939) describes the ovary of Mesopodopsis orientalis, Casanova (1977) that of Eucopia hanseni, Ishikawa and Oshima (1951) that of Neomysisjaponica, and that of PraunusJlexuosus, examined in this investigation, appears to be closely similar. The germinal area is located centrally and ventrally within the ovary and the eggs pass from this region into the ventral lateral areas where they

SP

hD

I

h

I

vd

sv

tc

C

SP

\

a

sv

3 spT$ .:,..... ..,.:.. ...:;,,;.. . ":. ...

'tc

I

D

sv

SSI

E

F

G FIG.11. Reproductive systems of mysids. A, Praunus fEexuosw, female system; B, P. f E e x w m s , male system; C, lateral view of male system; D, structure of male system of species of Archaemysis and Neomysis; E, cross section of testicular structures shown in D, arrow indicates movement of spermatozoa; F, spermatozoon of Mysis oculuta; G , spermatozoon of Praunus Jezuo8u.s (A, B. G from Mauchline, unpublished; C, after Labat, 1961; D, E after Kasaoka, 1974; F, after Fain-Maurel er al., 1975a). a, ejaculatory duct; h, heart; hp, digestive diverticulae; i, intestine; 1, larvae within marsupium; 0,ovary; ov, oviduct; p, penis; sp, spermatidic pouch; ssl, and ss2 first and second spermatocytic sacs; st, stomach; sv, seminal vesicle; tc, testicular cord; vd, vas deferens.

42

THE BIOLOGY OF MYSIDS

grow in size. They later move t o the dorsal lateral regions of the ovary and so posteriorly to the mouths of the oviducts. Nair briefly describes the histology of the ovary and reviews earlier work. The development of the eggs within the ovary of Metamysidopsis elongata was observed by Clutter and Theilacker (1971). They found that a period of about one week was required for eggs in the dorsal lateral regions of the ovary t o become invested with yolk. The testes of the male are located in the same general region of the thorax as the ovary in the female (Fig. 11). The form of the male system has been examined in Praunus jlexuosus by Labat (1961) and in Arch,aeomysis grebnitzkii and Neomysis awatschensis by Kasaoka (1974); both these authors review the earlier work. Holmquist (1959b), for Mysis relicta, and Kasaoka (1974) describe the histological structure of the system in considerable detail and reference should be made to their plates. The terminology adopted in this description is that of Kasaoka. The testicular cords, positioned ventrally, contain the spermatogonial cells. These cells produce the spermatocytes which, during maturation, cause the epithelial layer around the cords t o be formed into pockets, the spermatocytic sacs. These are not so obvious in Praunus jlexuosus as in the species investigated by Kasaoka. The spermatocytes move dorsally from the sacs to the spermatidic pouches. The wall of each pouch has a distinctive histological structure that includes a middle layer of muscle tissue. The spermatids develop within the pouches and mature spermatozoa pass through the relatively narrow duct into the seminal vesicle. Kasaoka describes this as a U-shaped tube while Labat shows it to be two tubes, a left and a right. A re-examination of P . jlexuosw during the present investigation (Fig. 11) confirmed Labat’s description of the system in P. jlexuosus. Kasaoka found spermatocytes in early phases of meiotic prophase I in the testicular sacs in both Archaeomysis grebnitzkii and Neomysis awatschensis. This is contrary to the findings of earlier authors. Holmquist ( 195913) suggested that little, if any, spermatogonial division occurred in adult mysids but that this must take place in young immature animals. Labat (1962) drew the same conclusion from his work with Praunus jlexuosus. The non-motile spermatozoa of the following species have been examined; Mgsis oculata by Retzius (1909), M . reEicta by Holmquist (1959b), Praunus jlexuosus by Labat (1962), Archueomysis grebnitzkii and Neomysis awatschensis by Kasaoka (1 974) and Praunus inermis in considerable detail by Fain-Maurel et al. (1975a, b). They are of t h e general form found in peracaridan crustaceans (Fig. 1l ) , consisting of a long narrow head and a tail or lash which is banded in structure.

3. THE

LARVAE AND REPRODUCTION

43

Reger et al. (1970) and Reger and Fain-Maurel (1973) have examined the origin and distribu.tion of the extracellular tubules surrounding the bundles of spermatozoa in the vas deferens. They are secreted by the Golgi complex in cells surrounding the developing spermatids but their function is unknown. Mature spermatozoa can often be obtained easily for examination from the vasa deferentia because species with well-developed penes frequently have spermatozoa stored there and in the ejaculatory duct (Fig. 11). An androgenic gland is associated with the outer wall of each ejaculatoryduct, occurring laterally. The gland is pyramidal to ovoid in shape, measures 50-200 pm in height, and has been described in Paramysis nouveli, Praunus Jlexuosus and Spelaeomysis longipes (Juchault, 1963 ; Meusy, 1963; Chaxniaux-Cotton et al., 1966; Nath et at., 1972). The chromosomes of several species have been examined by making squash preparations of the testes and early embryonic stages. Nair (1939) counted a haploid number ( n )as 22 in Mesopodopsis orientalis and Suomalainen (1954) recorded n as 18 in Neomysis integer but suggested that might be an under-estimate. Holmquist (1959b) re-examined N . integer and estimated n as 24-29. Anchialina agilis was the only species in which she made an accurate count of n ; it was 8. In all other species, the chromosomes stuck together a t meiosis and counting was usually impossible. Mitotic divisions within the embryos are less difficult to interpret but even here clear enough separation of individual chromosomes, to ensure accurate counting, was not found. Holmquist estimated 2n t o be 43-48 in Gastrosaccus spinifer, about 100 in Mysis relicta, 90-94 in Praunus jlexuosus, and 90-100 in Mysis litoralis, M . mixta and M . oculata. Later, Holmquist (1973, 1975) found n as 32 in Neomysis intermedia and probably 32 in N . mercedis while Archaeomysis grebnitzkii had a value for 2n of 10. This latter species, however, had a further smaller chromosome present in many of the preparations but its significance could not be determined. Labat (1959) determined the numbers of chromosomes in three species. He found n to be 48 in PraunusJlexuosus, so confirming Holmquist’s estimate. His value of n as 30 in Neomysis integer compares with an estimate of 24-29 by Holmquist. Paramysis nouveli has n as 56. Mating usually takes place a t night and lasts for only a few seconds. Observed mating positions are shown in Fig. 12. The possession of penes allows implantation of the sperm mass within the marsupium, but many genera do not have these organs, the vma deferentia opening at small papillae. Nouvel (1937) suggests that Praunus Jlexuosus discharges the spermatozoa into the water outside the marsupium and &at they are transported to the eggs by the water currents generated

44

THE BIOLOGY OF MYSIDS

FIG.12. Mating in mysids. A, Praunusflexuosus. Male approaches female in upper figure and moves into the coupling position in the lower figure (after Nouvel, 1937). B, Paramysis nouweli. Male approaches female in upper figure and moves into the coupling position in the lower figure (after Labat, 1954). C, Heteromysis armoricana coupling position (after Nouvel, 1940). D, Mesopodopsis orientalis coupling position (after Nair, 1939).

by the thoracic legs. Kinne ( 1 955), however, disagrees with Nouvel, discussing various possible uses of the male pleopods of Neomysis integer including the possibility that they actually transfer the sperm mass to the marsupium. The antennae may also be used as shown in Pig. 12A to hold the male against the marsupium; in the position shown in the lower figure, the sperm would be ejected when the male straightens his body. The lamellae of the marsupium are capable of muscular lateral movements, used for irrigation of the embryos. Such movement of an empty marsupium would probably be effective in drawing in the spermatozoa from the surrounding water. According to Clutter (1969) and Clutter and Theilackes (1971), male Metamysidopsis elongata transfer the spermatozoa to the empty marsupium of newly moulted females and the eggs are subsequently laid.

3.

THE LARVAE AND REPRODUCTION

45

Eggs sensu stricto do not occur in the marsupium since they are fertilized immediately they are extruded from the oviducts. The early embryos, however, are spherical or sub-spherical (Fig. 17) and their size is closely similar to that of ripe eggs within the oviducts. The term “early embryo” (Stage I larva) in the following discussion refers to this egg-like embryo, referred to as “egg” in Fig. 14 and Table 111. The number of early embryos contained in the marsupium of a female depends upon her body size, the size of the individual eggs and, in temperate and high latitudes, the season of the year. A relationship between the increasing body size of the female and increasing numbers of embryos in the brood is difficult to demonstrate in species of body length less than 10 mm and with small numbers of embryos, less than 15, in the brood. This relationship, however, has been shown to exist in the following species by the following authors : Anchialina agilis by Macquart-Moulin (1965); Erythrops elegans and E . serrata by Mauchline (1968, 1973b); Gastrosaccw lobatus by Macquart-Moulin (1965); G. psammodytes by Brown and Talbot (1972); G. spinifer by Mauchline (1971e, 197313); G. vulgaris by Matsudaira et al. (1952); Leptomysis gracilis by Mauchline (1969a) and Wittmann (1978); Mesopodopsis slabberi by Macquart-Moulin ( 1965); Metamysidopsis elongata by Clutter and Theilacker (1971); Mysidium columbiae and M . integrum by Brattegard (1973); Mysidopsis almyra by Price and Vodopich (19 79); M . angusta and M . gibbosa by Mauchline (1970b, 1973b); M . bahia by Price (1978); M . taironana, M . tortonesei and M . velifera by Brattegard (1973); Mysis mixta by Apstein (1906) and Mauchline (197313); M . relicta by Hakala (1978); M . stenolepis by Amaratunga and Corey (1975); Neomysis americana by Pezzack and Corey (1979); N . merc edis (awutschensis) by Heubach (1969); N . integer by Kinne (1955), Wi ktor (1961), Mauchline (1971b, 1973b) and Parker and West (1979); N . intermedia by Murano (1964a); N . japonica by Ishikawa and Oshima (1951); Paramysis arenosa by Liao (1951) and Mauchline (1971a, 1973b); P. intermedia and P. ullskyi by Boroditch and Havlena (1973); Praunus Jlexuosus, P. inermis and P. neglectus by Blegvad (1922), Liao (1951) and Mauchline (1965, 1971c, 1973b); Schistomysis kervillei by Mauchline (1971d, 1973b); S. spiritus by Liao (1951) and Mauchline (1967, 1973b); Siriella armata by Liao (1951); Taphromysis bowrnani by Beck (1977). This relationship has been examined in only one bathypelagic species, Eucopia unguiculata ( E . hanseni) ; Casanova (1977) found that this species had 8-23 embryos in the marsupium but the number was not related to the size of the female. Brood size can vary seasonally as illustrated in Fig. 13. The mean numbers of young per brood of ovigerous females of three different

46

THE BIOLOGY OF MYSTDS

size classes of Praunus inermis were determined as frequently as possible throughout the year in the Clyde Sea Area. The largest broods were produced during the early summer months. A similar analysis of the same size group of ovigerous Schistomysis spiritus was made in two populations; the first in the Clyde sea area, the second in Loch Ewe, north-western Scotland. The brood size changed seasonally in both populations but broods tended to be larger in the Clyde than in Loch Ewe. The range of body size, as indicated by the mean body length in Fig. 13, of ovigerous females within a population can also change seasonally and, of course, also influences the brood sizes produced by the total populations. The late summer and autumn breeding females are usually smaller in body size than those breeding in the spring and earij summer. Mauchline (1965-1971) has shown that a regular seasonal change in brood size takes place in Scottish populations of the following species, in addition to Praunus inermis and Schistomysis spiritus : Erythrops serrata, Leptomysis gracilis, Praunus Jexuosus, P. neglectus, Schistomysis kervillei and S . ornata. The work of Blegvad (1922) on Praunus spp. and Wiktor (1961)) Kinne (1955) and Parker and West (1979) on Neomysis integer also demonstrated a seasonal change. Wittmann (1978) found seasonal variation in the brood size of Leptomysis lingvura in the northern Adriatic Sea. Matsudaira et al. (1952) found such a change in populations of Castrosaccus vulgaris in north-eastern Japan but Brown and Talbot (1972) found no such change in South African populations of G. psammodytes. TWO species of Paramysis living in reservoirs of the Volga River, P . intermedia and P . ullskyi, showed marked seasonal changes in brood size (Boroditch and Havlena, 1973) as did populations of Neomysis mercedis (awatschensis) in the Sacramento-San Joaquin River estuary in California (Heubach, 1969), and N . intermedia in Japan (Murano, 1964s). The size of the earliest embryos in the marsupium can also vary seasonally in some species. Mauchline (1973b) found that the winter embryos were larger than the spring and summer ones in Leptomysis gracilis, L. lingvura, Neomysis integer, Paramysis arenosa and Praunus jlexuosus. No measurable difference was found seasonally in the embryos of eight other species examined. The smaller spring and summer embryos appeared to have fewer and smaller oil globules present in them. So far, we have been discussing variations in brood size within a species. Mauchline (1973b), however, has made a comparative study of embryo size and number, brood volume and the respective body sizes of the ovigerous females of many species. More data are now available and are summarized in Table 111. Casanova (1977) examined

Schistomysis spiritus

10

L

I

I

J

F

I

I

M

A

I M

I

I

I

J

J

A

I S

I

I

I

O

N

D

FIG.13. Praunus inermis. The seasonal change in the mean numbers of young present in the marsupium of females of A, 13-14, B, 12-13 and C, 11-12 mm total body length. Schistomysis spiritus. Upper figure: seasonal change in the mean numbers of young present in the marsupium of females nf 12-13 mm total body length in Kames Bay (dots) and Loch Ewe (triangles). Lower figure: seasonal changes in the mean body length of ovigerous females (broken lines) and mean numbers of young per female (solid lines) in Kames Bay (dots) and Loch Ewe (triangles] (after Mauchline, 1965, 1967).

TABLE111. BODYLENGTH, BROODSIZE AND SIZEOF THE “EGG” (STAGE 1 LARVA) IN MYSIDS. The bathymetric habitat of the species is indicated as: FW, freshwater; E, estuarine; S, shore; C, coastal; EP, epipelagic; MP, mesopelagic; BP, bathypelagic.

Species

Body Maximum length brood (mm) size

Range in body length

Acanthomysis sculpta A. strauchi A f r m y s i s guinensia Amathimyais cherados A. gibba

12 14 8.5 2.7 2.8

46 17 8 7 9

10-14 10-18 1.s2.7 24-24

A. polita Amblyops abbreviata Anchialina agilis

2.9 15 8.4 7.7 7 5.6 3.5 33 3.5 3.2 2.8 4 28 17 10.4 9 10

8 29 17 33 9 18 3 23 4 4 4 4 29 3 28 30 25

2.0-2.9 10-15 6-9 4.5-7.7 6.7-7.6 3.9-5.6 3.5-4.0 31-77 3.0-3.5 2.1-2.8 3.5-4.0 20-28 8 ~ 0 . 4 9-10

A. oculccta A. typica Aniaomysis hanseni Antarctomysis maxima Antromysis anophdinae Antromysis bahamensia A. cubanica Boreomysis arcticu B. megalops Bowmaniella bacescui B. bradie& B. johmoni

-

-

Range in brood size

“Egg” diamter

5-46 -

-

2-7 6-9 2-8 11-17 7-33

-

2-18 2-3 7-> 23

-

4 2-4 3-4 3-29 18-28 -

(mm)

-

0.3 0.3 (0.28-0.32) 0.27 0.4 -

1.3 0.25 0.25 0 0.8 -

-

Depth zone

Reference

S E S S S

Green (1970) Bacescu (1935) Bacescu (196th) Brattegard (1974a) Brattegard (1973, 1974a)

C BP C C C C C MP S S S S MP MP S C C

Brattegard (1974a, b) . Wigley and Burns (1971) Mauchline (1973b) Macquart-Moulin (1965) Ariani and Spagnuola (1976) Nouvel (1971) Nouvel (1967) Mauchline (1973b) W. M. Tattersall (1951) Brattegard (1969) Brattegard (1973, 1974a, b) Bacescu and Orghidan ( 1971) Jepsen (1965) Hoenigman (1963) Brattegard (1974b) Bacescu (1968d) W. M. Tattersall (1937)

B. p o r t o ~ c e w k B. sewelti Brasilornysis castroi

Ceratolepis hanzatu Cubanomysis jimenesi

Diamysis bahirens i s

D. frontieri D . pengoi Erythrops elegans E . erythrophtha1rn.a

E . parva E . serrata Eucopia, grimaldii E . sculpticauda Gaatrosaccus lobatus G. m a n g i i G. psammodytes ci. sanctus C . spinifer G. vulgaris Cnathophausia giyus G. gracilis

10 8.2 9 9.1

30 11

74-10

10 16

7-9 5.6-9.1

6-10 4-16

14 4 3.2

22 3 4

13-14 3.5-4.5 2.2-3.4

2-5 2-6

6 11.5 5 13 5.9 7-1 4.5 3.5

7 28 7 20 13 15 8 4

6.5-8.0 S11.5 8-13 4-6 5.7-7.1 3.2-3.5

8.7 32 35 10.4 9.5 15 8 12.7 17 127 76

27 8 1 42 35 69 12 67 100 9 69

-

6.5-10 27-55 30-60 74-10.4 7.5-8.5 13-2 1 (23

10-14 14-18 49- > 100

1-30

-

3-7 15-28 2-7 8-20 1-13 1-15 2-4 2-27 4-8 < 20-42 12-69 6-12 21-67 30-106 -

2-97

0.38 (0.36-0-4) 1.0

-

0.25 (0.24-0.27) 0.31 0.38 0-4 0.29 (0.26-0.30) 0.35 1.5 0.5 0.46 (0.54-0.59) 3.2

C C

C C

EP S S

Wigley and Burns (1971) Brattegard (1974a) Bacescu (1968a) Brattegard (1973, 1974a) Fage (1941) Bacescu (1968~) Brattegard (1973, 1974a, b)

C Genovese (1956) E Bacescu (1954) C Nouvel (1965) E Bacescu (1954) C Mauchline (197313) MP Wigley and Burns (1971) M P Macquart-Moulin (1965) C Brattegard (1974a) C BP BP

C-

s-c

S C S S BP BP

Mauchline (1973b) Fage (1942) Fage (1942) Macquart-MouKn (1965) Bacescu (1975) Brown and Talbot (1972) Hoenigman (1964a) Mauchline (1973b) Matsudaira et al. (1952) Mauchline (197313) Fage (1941);Mauchline (197313) cowt.

TABLEI I I - c o n t .

Species

Body Maximum length brood (mm) size

Range in body length

140

350

G . longispina C. zoea

42 80

1 42

40-64 40-105

Hemimysis anomala H . bmornae

11 8.7 8 6 4 4 8 4.5 6 7 7 7 5 4.5 10 9 7.2 13.9 11.9 11.3

31 35 9 20 4 7 15 5 2 5 5 5 4 8 23 15 3 76 26 51

8.5-1 1 7-1 1 6.5-8 6-8 3.3-4

G . ingens

Heteromysis urmoricana H . bermudensis H . elegans H . formosa H . guitarti H . gymnura H . harpax H . mariani H . rubrocincta H . zeybnica Idiomysis tsumzantali Katamysis wurpaehowskyi Leptomyaia apiops

L. gracilis L. lingvura

130-> 145

-

5-8 4-4.5 4-6 6.5-7.8 6-7.5 6.5-8 -

4’2-4.5 5-10 9-13 6-8 9-16 8.5-12 7-1 1.5

Range in brood size

“Egg” diameter (mm)

4.3-200

8-3 1 5-35 4-9 9-20 4

-

13-15 3-5

-

1-5 3-5 2-5 2-4 5-8 3-23

-

(0-46-0.54)

Reference

Mauchline (1973b); Childress and Price (1978) BP Fage (1941); W. D. Clarke BP Fage (1941); Mauchline (1973b) C Bacescu (1954) Mauchlhe (1973b) C Bacescu (193613) C Nouvel (1940) C Brattegard (1973) C C Brattegard (1974a) C Wigley and Burns (1971) C Bacescu (1968b) S Fage and Monod (1936) S 0. S. Tattersall (1962) C Bacescu (1970b) C Bacescu (1968b) C W. M. Tattersall (1922) S Bacescu (1973a) E Bacescu (1954) C Bacescu (1966) C Macquart-Moulin (1965) C Mauchline (197313) C Macquart-Moulin (1965) C Mauchline (1973b) BP

20-42

3-76 13-26 7-51

Depth zone

L. mditerranea L. pereai Linznmysis benedeii i Lophogaster afinis L. typicw

-

9-27

-

6-8

-

18 8.5 15 24 23

27 8 40 25 34

8-8.5 7-15 17-30 17-23

Mesacanthmysis pygmaea 2.5 Mesopodopsis orientalis 7.6 M. slabbe& 11 M . zeylanica 5.6 Metamy&p&s elongata 6.5

5 9 20 4 30

7-8 7-1 1 6'2-5.6 4-7

3-5 8-10 5-20 1-4 4-30

0.32 0.35

M . insularis M. s ~ f t i

13 20 14

13 13-20 5-14 3-8

0.4 0.25

8

4.5-5.0 5.5-6.5 4-5 3-4-3.9

3.2 23 7 7

6 10 11 9

3.0-3.2 22-23 5-7 4.8-7

5-6 1-11 2-9

M. angwta M . ankeli

5-6 5.9 4-8 6.8 3.4

10 6 13 17 10

3-6.6 5.1-6.5 4.3-5.8 5-7 2.6-3-4

2-10 3-6 3-13 5-17 5-10

M . arenoaa

4.3

4

-

-

Mysidella minuta Mysidetes posthon Mysidium colunzbiaa M . integrum MYsidOpSi8 almyra

5.0

7 5 3.9

-

20-40 22-25 22-34

0.8

-

-

0.25 (0.20-0.26) 0.34

0.40 (0.40-0'44) (0.40-0.42)

-

(0.35-0.39) 0.29 (0.28-0.30) -

C

Liao (1951) Bacescu (1966) E Bacescu ( 1954) MP Fage (1942) MP Fage (1942) ; Hoenigman (1963) S Nouvel (1967) E Nair (1939) S Macquart-Moulin (1965) S Nouvel (1954) S Clutter and Theilacker (1971) S Quintero and Roa (1973) C Bacescu (1969) C Brattegard (1970a) S Brattegard (1973, 1974a)

C

-

C MP S S

Brattegard (19744 0. S. Tattersall (1965) Davis (1966) Brattegard (1973, 1974a, b)

S S S S

Brattegard (1973, 1974a) Price and Vodopich (1979) Price and Vodopich (1979) Mauchline (1973b) Brattegard (1973, 1974a)

S

Brattegard (1974b)

C

cont.

TABLEI I I - c o n t .

Species

M . bahia

Body Maximum length brood (mm) size

Range in brood size

“Egg” diameter

(mm)

Depth zone

-

0.25

4-6 4.8-5-0 2.8-4-4 2.1-3.0 3-6-6.2

3-12 2-4 12-16 9-16 17-60 5-34 8-10 2-4 2-8 3-1 1

11 8 21 97

2.6-4.4 4.2-6.2 29-35 13-23

3-1 1 2-8 12-2 1 9-97

0.28 0.34 (0.32-0’36) 0.34 0.40 1* 7 6-1 a92 0.60-0.64

21

40

13-21

10-40

0.66

Fw

22

32

15-24

5-32

-

Fw

188 245

-

46-1 88 140-245

0.4

-

C C

26

6-16

626

0.4

C

5.0 5.3 3.0 6.5 7.5 4.5 14-5 12.5 5.8

8

4.4 2.8 6.2

14 4 23 16 16 60 10 34 10 4 8 11

M . velifera M . virgulata My& litoralis M . mixta

4.4 6.2 35 22

M . relicta

M . bispinulata M . camelina M . coralicolu M . cultl.ccta M . dide~!phys M . eremita M . gibbasa M . mawhlinei Id.mortemeni M . taironana M . tortonesei

Range i n body length

5.0

M . stenolepis

25.5

Neomyeis amrkxzna

12

4.0-6.6 2.1-3.0

-

7’5-8.5 4.0-4.5 10-15

-

-

-

0.47

-

0.36 0.35

-

E E S S S S C C

C S S S S S C C C

Reference

Nimmo et al. (1978) Price (1978) Brattegard (1974a) 0. S. Tattersall (1955) Bacescu (1975) Brattegard (1973) Mauchline (1973b) 0. S. Tattersall (1962) Mauchline (197313) Brattegard (1974b) Brattegard (1973) Brattegard (1973, 1974a, b ) Brettegard (1973, 1974a) Brattegard (1973, 1974a, b) Brattegard (1974a) Mauchline (197313) Apstein (1906); Mauchline (197313) Tattersall and Tattersall (1951) Reynolds and De Graeve (1972) Wigley and Burns (1971) Amaratunga and Corey (1975) Wigley and Burns (1971); Mauchline (1973b)

N. rnercedia (awatschensis) 17 N. integer 16.6 16 N. intermedia 16 N. japonica 13 Paramysis arenosa 9.2 10 P. bacescoi 9 P. i n t e d i a 15 14

57 72 55 46 30 33 34 15 42 36

P. kroyeri P. nouveli P. pontica P. ullskyi

16 10 19 18 21

Praunua jlexuosus

25.3 23 24 25 13.9 16 15 21.3 21 4.6 10.9 11.8 11.5 15

P. inermis P . neglectua Prornysis atlccntica P s e u d m afine P. craa&entatum P. ear& P. scholhrtenais

7-17

-

10-17

5-57 15-72

10-16 12-16

8-55 1-46

5-9.5 5-10 5.5-8.5 8-15 6-14

2-33 4-34 4-15 7-42 7-36

(0'45-0.60)

30 28 43 30 62

10-16 6.5-10 10.5-1 9 13-18 12-2 1

6-30 6-30 6-43 6-30 17-62

(0.6-0.8)

63 76 40 44 56 50 > 28 81 > 42 5 11 23 14 18

13-24 12-23 17-24 16-28 10-16 9-16 9-15 14-23 13-2 1 3.6-4.6 8-11 10.9-11.8 -

4-63 17-76 14-40

(0.78-0.8 1) 0.67 -

-

-

15-56 4-50 5->28 13-81 5-> 42 2-5 1-1 1 15-23 -

(0.5-0.54)

-

0.50 (0.47-0.49)

-

0.80

-

(0.32-0.36) 0.41

-

E E E FW E S S S C

C

E S C C C S S S S S S S S S C C BP BP E

Heubach (1969) Mauchline (1973b) Kinne (1955) Murano (1964a, b ) Nakazawa (1910) Mauchline (1973b) Liao (1951) Labat (1957) Bacescu (1954) Boroditch and Havlena (1973) Bacescu (1954) Labat (1957) Bacescu (1954) Bacescu (1954) Boroditch and Havlena (1973) Mauchline (1973b) Liao (1951) Blegvad (1922) Wigley and Burns (1971) Mauchline (1973b) Liao (1951) Blegvad (1922) Mauchline (197310) Blegvad (1922) Brattegard (1973, 1974a) Wigley and Burns (1971) Murano (1974b) Murano (197413) 0. S. Tattersall (1955)

TABLE111-cont. ~~

Species

Body Maximum length brood (mm) size 8

12

14.8 14.3 14.5 13.5 23 24 11 9.5

47 23 29 30 48 99 15 13

S. clausii

10 8

S. jaltensis

Range in body length

Range in brood size

“Egg” diameter (mm)

Depth zone

Reference

2-12

S

8-16 11-16 9.5-1595 9-14.5 19-23 16-24 10.5-1 1.5 7.8-9.5

4-47 2-23 4-29 3-30 31-48 30-99 6-13

S C S S S S C C

Mauchline (197313) Mauchline (197310) Mauchline (197313) Liao (1951) Mauchline (197313) Liao (1951) Ariani and Spagnuolo (1976) Brattegard (1974a)

23 12

5-9 6-8

10-23 5-12

C C

8-1 11.9 8.5

16 22 13

5-8.5 10-11.9 8-13

8-16 -

C C C

Speluewnysis longipes

18.5 9 8.6 5

46 7 6 9

14-18.5 5-9 5-8.6 4.4-5.1

2-7 2-6 7-9

Taphromyeis b o m n i Tenagomy& macropsis

10 9

17 25

7-10 -

6-17 -

Mauchline (197313) Genovese (1956);Ariani end Spagnuola (1976) Macquart-Moulin (1965) Mauchline (1973b) Macquart-Moulin (1965) ; Ariani and Spagnuola (1976) Hoenigman ( 19634 Bacescu and Mayer (1961) Hoenigman (1963) Pillai and Mariamma (1964); Nath (1973) Beck (1977) Barry (1956)

Rhopalophthalmus longicauda Schistomysis kervillei S . ornata S. spiritus Siriella armata

S. castellabatensis S. chierchiae

S. norvegica S. thompsonii

7.6-8

-

C

E E S S C

0. S. Tattersall (1957)

3.

55

THE LARVAE AND REPRODUCTION

the brood sizes of Eucopia unguiculata ( E . hanseni) but measured carapace length as’an index of total body length; his data, are not included in Table 111. Larger sized early embryos tend to be produced by larger species but there is considerable variation (Fig. 14). There appears t o be a minimum size of embryo, about 0.25-0.30 mm in diameter. These embryos are produced by species of less than 5 mm total body length with an exception, on present information, of Metamysidopsis swifti (Table 111). Another species that has early embryos

10

102

103

104 Body length3

105

106

Fio. 14. The relationship of ‘‘egg” (Stage I larva) volume to total body length (raised to the third power) of the parent mysid. Dots are coastal and epipelagic species, triangles are meso- and ‘bathypelagicspecies. The regressi6n equations for the lines and their respective correlation coefficients are : coastal, epipelagic: log “egg” volume = 0.4510 log L8--2*6117,T = 0.8197*** meso-, bathypelagic: log “egg” volume = 0.9367 log Ls-4.2395, T = 0.9577***

A 100

I-

I

ee

/

A

'

ee

A

1 10

I

I

I

I

1

lo2

lo3

lo4

lo5

106

Body length3 FIG.15. The relationship between maximum numbers of young per brood and total body length (raised to the third power) of the parent mysid. Dots are coaatal and epipelagic species, triangles are meso- and bethypelagic species. T h e regression equations for the lines and their respective correlation coefficients are: Coastal, epipelagic: log number of young = 0.4264log L3+0.0281,r = 0.8107*** Mew-. bethypelagic: log number of young = 0.1547 log Ls+0.5667, r = 0.3631

3.

THE LARVAE AND REPRODUCTION

57

that are markedly smaller relative tc its body size is Mysis stenolepis. Meso- and bathypelagic species tend t o produce eggs that are larger relative to their body size than epipelagic and coastal species (Fig. 14). The numbers of young per brood varies greatly between species and sometimes between closely related species. There is, nevertheless, a tendency for larger species t o have larger numbers of young per brood (Fig. 15). The species, however, are divided into two natural groups. The first consists of epipelagic and coastal species while the second consists of the meso- and bathypelagic species. These latter species (Fig. 15) produce smaller numbers of young per brood relative to their body size than do the epipelagic and coastal species. Brood volume is directly related to body size (Fig. 16). The volume of the brood is obtained by calculating the volume of the spherical or near spherical early embryos and multiplying by the number of embryos present. No distinction exists between the deep living species and the others ; the few data suggest the possibility that the largest mesoand bathypelagic species may produce slightly larger broods relative to their body size than the other species (Fig. 16). The embryology of Hemimysis lamornae, Mesopodopsis orientalis, Neomysis integer and Praunus jlexuosus has been studied in considerable detail (Nusbaum, 1887 ; Wagner, 1896; Nusbaum and Schreiber, 1898; Manton, 192813; Needham, 1937; Nair, 1939). None of these papers has been superseded, nor is it possible to summarize them adequately here. Consequently, direct reference should be made to these descriptions which include many plates. The larval development of mysids takes place entirely within the marsupium. Detailed descriptions of the histology of successive larval stages are given by Nusbaum (1887) for Praunus jlexuosus, by Wagner (1896) for Neomysis integer, by Manton (1928b) for Hemimysis lamornae and by Nair (1939) for Mesopodopsis orientalis. The external morphology of the larvae of the following species has been described : Acanthopysis sculpta by Green ( 1 970) ; Boreomysis arctica by Jepsen (1965) ; Gastrosaccus psammodytes by Brown and Talbot (1972) ; G. vulgaris by Marsudaira et al. (1952); Gnathophausia species by Fage (1941); Metamysidopsis insularis by Quintero md itoa (1973) ; Mysidium columbiae by Davis (1966, 1968) ; Mysis relicta by Berrill (1969) ; Neomysis integer by Kinne (1955); N . intermedia by Murano (1964b); Schistomysis spiritus by Almeida Prado (1966). The development within the marsupium can be divided into three stages (Nair, 1939; Matsudaira et al., 1952; Jepsen, 1965; Davis, 1966, 1968) which correspond with the stages described by Mauchline (1972, 1973b) as “eggs”, eyeless larvae and eyed larvae.

A

/

/* 10

. I

I

1o2

1o3

1 1o4 Body length3

I

1

1o5

106

FIG.16. The relationship between volume of the brood and total body length (raised to the third power) of the parent mysid. Dots are coastal and epipelagic species, triangles are meso- and bathypelagic species. The equation for this relationship and it8 correlation coffioient are : log brood volume = 0.9147 log L3--2.6280, r = 0.9583***.

3.

THE LARVAE AND REPRODUCTION

59

Stage I . The early embryo, at first egg-like (Fig. 17A) but later with rudiments of antennae and abdomen developing (Fig. 17B). It is still within the egg membrane which is shed at the end of this stage. Stage I I . The larva has hatched from the egg membrane by puncturing it with the abdomen (Fig. 17GF). The antennae and thoracic appendages develop during this stage and the eyes become.pigmented. This stage terminates in a moult.

A

0

C

Fro. 17. Larvae of a mysid. A, egg-shaped embryo (“egg”);B, embryo with developing abdomen; C-F, stage I1 larvae at successive stages of development; G , stage I11 larva after the first larval moult and with the eyes stalked; H, stage I1 larva of a Cfnathophausiaspecies. Oil or yolk is represented by areas of darker stippling.

Stage I I I . The moulted larva now has the eyes on stalks (Fig. 17G). No lith is present in the developing vesicle of the statocyst in the uropods. This stage also terminates in a moult that takes place as, or shortly after, the larvae are released from the marsupium. Davis (1966) found that the early Stage I embryos (Fig. 17A) of Mysidiurn columbiae swelled within the marsupium and burst a thin

60

THE BIOLOGY OF MYSIDS

external membrane from which they emerged. This membrane is additional to the one shed a t the end of Stage I but no other authors have reported its presence. The embryos of most species have globules of oil (yolk) present within their tissues. These are usually in the form of many small globules but Schistomysis spiritus also has a large single globule present in the central region of the Stage I larva and in the anterior region of the Stage I1 larva, These globules are spread throughout most of the body of Stage I and I1 larvae but, as they decrease in volume relative to the body volume of the larva, they become concentrated in the anterior dorsal regions by the end of Stage 11 (Fig. 17F). The quantity of oil varies from species to species and its presence is very marked in larvae of the bathypelagic Gnathophausia species (Fig. 17H). Herring (1974) found that the large eggs of pelagic decapods have relatively high lipid contents and a low density. Their range of density was 1~091-1.025g/ml which compared to the value of 1.04 g/ml for the Gnsity of the Stage I larvae of Eucopia unguiculata. All the p v a e within a single marsupium are at the same stage of developmeht. They are regularIy orientated, their heads pointing posteriorly (Fig. 11A, p. 40) and closely packed together. Occasionally a Stage I1 larva occurs among Stage 111 larvae but this is rare and probably results from adoption (p. 62). Mauchline (1973b) found indications of a rate of mortality of about 10% during marsupial development. Amaratunga and Corey (1975) found fewer Stage I11 larvae in marsupia than Stage I indicating a loss of larvae in Mysis stenolepsis. The Stage I larva of Praunus jlexuosus, Mysis relicta and Neomysis americana is inactive and only morphological differentiation takes place. The Stage I1 larva exhibits a slow rhythmic beating of the heart and contractions of the gut; the gut contractions cause movement of the oil globules (yolk) (Berrill, 1969, 1971). The entire body of the late Stage I1 larva of Praunus jlexuosus and Neomysis americana flinches within its integument which has stretched and now serves as an enveloping membrane. Once this integument is shed the abdomen is free t o flex in the manner of the adult “escape reaction”. The appendages of these, now Stage 111, larvae have periods of active rhythmic movement appropriately described by Berrill as fluttering. The larvae have become increasingly active throughout Stage 111. They are markedly larger in size than the comparable Stage I embryos (Table IV and V) and the marsupium is distended to contain them. The brood lamellae are moved laterally in a rhythmic manner by the female to irrigate the larvae and consequently it would be surprising if some accidental loss of larvae from the marsupium did not occur.

3.

61

THE LARVAE AND REPRODUCTION

TABLEIV. LENGTH(MM) OF LARVAEWITHIN THE MARSUPIUMAND OF JUVENILES ON EMERGENCE OF EPIPELAUIC AND COASTALSPECIES( A f t e r Mauchline, 1972) Species Acanthomysis sculpta Anchialina agilis Eythrops elegans E . serrata Gaatrosaccua psammodytes G. spinifer G. vulgaris Hemimysis lam& Heteromysis formosa Leptomysis gracilis L. lingvura Mesopodopais orientalis Metamymdopsis elongata Mysidiurn columbiae Mysidopsis angusta M . didelphys M . gibboaa Myais mixta M . oculata M . stenolepis Neomyais americana N . integer N . intermedia Paramysia arenosa Praunus jexuosua P. inermk P. negbctua Paeudomma affgne Schistomysis kervillei S. ornata S. spiritua Siriellu armata S. clauaii S. jaltensis Taphromysis bowmani

Stage I

Stage 11

Stage 111

-

1-P1.6 0.9-1.1 0.8-0.85 0.8-0'9 1.6-1.8 1.1-1.3 1*o-1*2 1.0 1.25 1*3-1.4 0.7-0.9 0.5-0.7 0.8-1.1 0.8-0.9 1.3-1.5 0.9-1.1 1.3- 1.7 4.4-5.7 1.3- 1-6 0.6-1.2 1.3-1.5 1'0-1 '4 1.2 1.7-2.2 1 3- 1.7 1.7-2.1 1.3-1.5 1.5-1.7 1.1-1'3 1.3-1.4 1'2-1.3 0.9-1.0

1.8 1.3 1.4 2.0 1.7-2.1

-

0.38 0.35 0.5 0.45-0.50 0'54-0.59 0.5 0.4 0.38-0.48 0.40-0-55 0.35 0.4 0.4 0.35-0'42 0.4-0.5 0'35-0.40 0.6-0.64 1.76-1'92 0'49-0.55 0.41 0.5-0.6 0.5 0.4-0.5 0.7-0'9 0.6-0.8 0.7-0.9 0.4 0.6-0.7 0.6-0.7 0.5-0.6 0'5-0.6 0.45-0.5 0.5-0.6

O n emergence c. 2.0 2.0 2.5

-

-

1.P1.6 1.7 1.7-1.9 1.7- 1*8 1-3 1.2

1.6-1.9 1.2-1 '3 1.7-2.2 6.6-7.0 1.6-1.8 1.3 1-9-2.1 1.4-1.7 1-4-1.6 2.5-3.0 2.2-2.6 2.7

1.6-1.9 1.8-2.2 1.7-2.0 2.2-2.3 1.6

2.0

-

C. C.

2-5 2.5 1-5 -

2.5 1.5 2.0 2.0 1-5 2.5-3.0 2.5 2.5-3.0 C. 3.0 2.0 2.5 2.0 2.5 -

Female mysids not only protect the larvae by carrying them within a marsupium but have also evolved behavioural patterns that enable them to care for the brood in a much wider sense. Wittmann (1978) found that ovigerous Leptomysis lingvura are capable of catching larvae that have escaped from the marsupium. The process of catching them appears to be identical with that for catching larger particles of

62

THE BIOLOGY OF MYSIDS

TABLEV. LENGTH(M M ) OF LARVAEWITHIN THE MARSUPIUMAND OF JUVENILES ON EMERGENCE OF MESO- AND BATHYPELAGIC SPECIES (After Mauchline, 1972) Species Antarctomysis maxima Boreomyais arctica Erythrops erythrop-a Eucopia grimaldii E. sculpticauda Gnathophawia gigas G. gracilis G. ingens G. longispina G. zoea Lophogaster typicus

Stage I

Stage II

Stage III

1.3 0.3 0.4 1.5 -

3.8 1.0-1-2

5 1.4 4.0

3.2 4.0 3.4 0.8

7 5’0-9.1 13 6 10-12 2

O n emergence

22.4 13.3-16.0

13 5

food. The female, however, when the larva reaches the mouthparts usually does not eat it but’ moves it posteriorly by movements of the thoracic legs so that it enters the marsupium via the anterior or, occasionally, the ventral space between the brood lamellae. Adult males and immature males and females will also catch escaped larvae but they either release them immediately or eat them. Larvae that are at a later stage of development than those of the female’s own brood are adopted more frequently than younger larvae. Leptomysis lingvura adopts larvae of L. burgii, but less frequently than larvae of its own species. It can also distinguish the relative age and species of parts of larvae introduced to it and freely adopts parts of larvae of its own species. Adopted parts of larvae remain in the marsupium along with introduced sand grains so indicating that no active cleansing of the larvae takes place within the marsupium. Wittmann shows experimentally that adopted larvae survive and are liberated from the marsupium in the usual manner. Their liberation is usually before that of the female’s own brood because the adopted larvae are usually older. The female moults soon after liberating her own brood and so younger larvae that have been adopted and are not yet ready for release are lost a t this moult. Wittmann found that 0.8% of females of L. lingvura and 0.25% of females of L. burgii in natural populations in the northern Adriatic Sea had adopted one or more larvae; this was determined by the presence of one or more older larvae among broods. These adopted larvae represented 0*082%and 0.034% respectively of the total numbers of larvae within the broods of all the females of these two species examined.

3.

THE LARVAE AND REPRODUCTION

63

Wittmann’s observations have been confirmed by Mauchline and Webster (unpublished). The following species were found to be able to adopt escaped larvae : Mysidopsis gibbosa, Neomysis integer, Paramysis arenosa, Praunus JEexuosus, P. inermis, P . neglectus and Xchistomysis spiritus. Larvae that were older than those of the female’s own brood were ado ted more frequently. Dead Iarvae were usually rejected completely or ten but were not adopted. Pruunus jkxuosus and P. neglectus w o d adopt each other’s larvae and those of P. inermis; P. inermis, however, would not adopt the larvae of the other two species. Consequently, this adoption of free larvae decreases the potential mortality of larvae arising from their accidental loss from the marsupium. The duration of marsupial development is related to environmental temperature (Blegvad, 1922; Ishikawa and Oshima, 1951 ; Murano, 1964a). The duration in Praunus species ranges from 14 to 44 days depending upon the seasonal environmental temperatures in Denmark and in Scotland (Blegvad, 1922; Mauchline, 1965); the seasonal range of temperature is approximately 7-18°C. The duration of the development in Neomysis intermedia in Japan ranges from 6 to 9 days over a range of temperature of 21-3OoC, while that of AT.japonica is almost linearly related to temperature over the range 12.2-29.5OC (Table VI). Some examples of development times are given in Table VI. The cold water species, Boreornysis arctica, Mysis relicta and the bathypelagic Gnathophausia ingens have the longest reported development times, while the shortest is reported for Mesopodopsis orientalis at 25-29°C. Salinity affects the duration of the larval development of Neomysis integer. Vlasblom and Elgershuizen (1977) found that the range of salinity within which the larvae develop is more restricted than the total range in which the female can survive. Larvae adapted to a salinity of 7 x 0 had a development time of 15-18 days while those adapted to a salinity of 23% had a comparable development time of 17-22 days. A pair of tubes extend ventrally from the female’s thorax into the marsupium of species in the genus Neomysis. They are illustrated for N . integer by Vorstman (1951) but their function is unknown. McLusky and Heard (1971) found that Praunus jlexuosus can regulate the composition of the marsupial fluid hyper-/hypo-osmotically, with a notable degree of control. There was a linear correlation between the osmotic concentration of the blood and that of the marsupial fluid but the degree of regulation was much strider in the case of the blood. Consequently, larvae within the marsupium of species living in fluctuating salinities may be protected from the extreme ends of the range.

i

TABLEVI. THE DURATION (DAYS)OF

THE

DEVELOPMENT OF THE LARVAL STAGES WITHIN PERIOD O F M A R S U P I A L RESIDENCE

THE

MARSUPIUM AND

THE

TOTAL

Duration of Temperature

species Acanthmysis sculpta Boremysis arctica Gaatrosaccus vulgaris Gnathophawria ingens Hemimysis speluncola Mesopodopsis orientalis MetanZyaidopsis elongata Mysidium columbiae Mysis relicta Neomysis americana N . integer

N . japonica Praunwr jexuosus 0

("(3 12 V7) 10.9 3.5 12 14 23 25-29 17-19 3 2-10 0.2-3'8 12-15 10 16 15 12.2-29.5 12-15

Stage I

Stage I I

Stage I I I

Total

1

3

2

c. 28

c. 40

c. 30

c. 100

11 163-202

8 105-154 9-14 -

25 313-530 26-31} 17-19

6

6-7

-

2-3 2-3

-

10-11

-

I

5-6

11-13

4 10 6-7 c. 150 c. 270 c. 21 23-25 12-14 15-22 6-17* c. 21

Reference Green (1970) Jepsen (1965) Matsudaira et al. (1952) Childress and Price (1978) Macquart-Moulin and Patriti (1966); Gaudy and Guerh (1979) Nair (1939) Clutter and Theilacker (1971) Davis (1966) Lssenby and Langford (1972) Berrill (1971) Pezzack and Corey (1979) Pezzack and Corey (1979) Vlasblom and Elgershuizen (1977) Ishikawa and Oshima (1951) Berrill (1971)

Log incubation period = 1.9313-0.0559T where T is environmental temperature in 'C (Ishikawa and Oshima, 1951).

3.

THE LARVAE AND REPRODUCTION

65

Emergence of larvae from the marsupium generally takes place a t night and over a short period of time, a few minutes t o a n hour (Green, 1970; Berrill, 1971). The late Stage I11 larva within the marsupium is active and shows rhythmic twitching of the body (Berrill, 1969, 1971). The larvae are released from the marsupium when the female spreads the lamellae laterally and the larvae fall out, one by one (Nair, 1939). The larvae are unable t o swim until after the moult that terminates Stage I11 of development. Berrill (1971) found that this moult took place as the larvae were emerging from the marsupium. Nair (1939) suggests t h a t it takes place in Mesopodopsis orientalis within 10-20 min. of emergence, while Amaratunga and Corey (1975) found that the larvae of Mysis stenolepis moulted up to 3 h after emergence. This postmarsupial moult is synchronized in broods of Neomysis arnericana; Pezzack and Corey (1979) found that a t 4, 10, 16, 22 and 25°C it took place 312, 144, 72, 48 and 24 h respectively after emergence. It is likely, however, that this moult takes place irl the natural environment in most species very close t o the time of emergence. Immobile larvae are helpless and would be subject t o heavy predation. One possible source of such predation would be cannibalism by the female but Amaratunga and Corey (1975) did not observe any such behaviour in Mysis stenolepis. One notable feature of this post-emergence moult is the development of the lith in the vesicle of the statocyst, a feature absent in Stage I11 larvae. The young mysids are now miniature adults, usually 1.5-3mm in total body length, except in the large G n u € ~ o ~ ~ a u s i a species where they may attain lengths of' over 20 mm (Tables IV and V).

CHAPTER 4

VERTICAL DISTRIBUTION AND MIGRATION Mysids are predominantly marine organisms, a few species living in fresh water and in specialized environments such as caves and wells. Some 36 species live in non-marine habitats, about 160 are more or less restricted t o the littoral environment, about 300 t o the neritic region, 30 are epipelagic and over 200 live in the meso- and bathypelagic regions. Information on the vertical distribution of many species is not available. Some species, reported as meso- or bathypelagic, are probably not markedly pelagic in habit but usually restricted t o the hyperbenthic region-the water layer adjacent to the sea bottom. Nets mounted on sledges (Fig. 18) are especially useful in sampling mysids as found by Bossanyi (1957), Beyer (1958), Hesthagen (1973) and several others. Comprehensive sampling of mysids and investigation of their vertical distribution within a coastal area is difficult because various types of sampling equipment have to be employed. The pelagic species or hyperbenthic species that have migrated upwards can be sampled with the conventional plankton nets. These can range from the conical tow-nets to the Isaacs-Kidd Midwater Trawl which has been used by Pequegnat (1965) and Fried1 (1971b), or as modified by Pearcy et al. (1977) to sample offshore species. The Rectangular Midwater Trawl (RMT) and other offshore samplers used to capture euphausiids (Mauchline, 1980) are equally suitable for pelagic mysids. Littoral and estuarine species can often be taken with hand dipnets, push-nets, or very simply with the hand-pulled D-net shown in Fig. 18. A modification of the push-net that quantitatively samples contiguous parts of a transect is described by Miller (1973) and could be adapted for shallow water sampling of littoral mysids. The Ocklemann detritus sledge (Fig. 18) is heavier to use by hand than the D-net but is favoured by Brattegard. A further multilevel net system to that shown in Fig. 18 is described by Ellertsen (1977) and is potentially useful in littoral and sub-littoral regions. Heavier and more robust equipment is required to sample the neritic species. Various forms of beam trawl incorporating runners of 10-20 cm diameter are

4.

VERTICAL DISTRIBUTION AND MIORATION

67

A

FIG.18. Various towed samplers used to catch mysids. A, the D-net used by Mauchlhe; B, bottom plankton sampler designed by Colman end Segrove (1955) ;C, the modified Beyer epibsnthic sledge as described by Oug (1977); D, the Ockelmann detritussledge described by Brattegard (1973); E, the multilevel net described by Fager et al. (1966).

used to tow over soft substrates. One or several nets, in addition to the main trawl, can be attached to the wooden beams of such trawls to provide samples at heights of 1-2 m above the surface of the sea bottom. Several more specialized samplers are available (Fig. 18). Wickstead (1953), Clutter (1965), Macer (1967) and Omori (1969b) have designed closing epibenthic sledges that are all excellent samplers for mysids. Poirier et al. (1969) have modified the Macer sledge,

68

THE BIOLOGY O F MYSIDS

and the larger Beyer system (Fig. 18) is used by Hesthegan and Gjermundsen (1978) and Schnack (1978). Light lures can be incorporated in many of these nets as first used by Herdman (1890) and later by Brattegard (1970a) and Bacescu (1975). Mysids can avoid nets and can be observed escaping from samplers used in the littoral region. Fleminger and Clutter (1965) studied the escape of Metamysidopsis elongata and found that sampling was more efficient in darkness. They also discovered that less avoidance by individuals took place in dense populations than in populations that were dispersed. Quantitative sampling of mysids, many littoral species of which form aggregations, is extremely difficult. The difficulties are increased in the case of some Gastrosaccus and other species that partially or totally bury themselves in the sediment. during the daylight hours. The capture of shallow living species of mysids in good condition for experimental work is relatively easy and many can be maintained in the laboratory for considerable periods, of time. Greve (1975) has reared Praunus Jlexuosus and Mesopodopsis slabberi through a t least one generation in his “Meteor Planktonkuvette”. Deeper living species especially the meso- and bathypelagic species, are more difficult t o obtain in excellent condition. Childress et al. (1978) have solved the problem by designing a thermally protected cod-end for their nets. The terminal region of the net is contained within a heavy-walled (1-15 cm thick) polyvinylchloride tube with valves that can be closed. The tube is open during the shooting and towing of the net but it is closed for retrieval. The animals within the tube are then thermally insulated from the outside environment as the net is raised through the warmer surface layers of the ocean. Gnathophausia species caught in this manner have been in excellent condition for experimental investigations (Childress and co-workers). The depth of occurrence of the different species is indicated by Mauchline and Murano (1977) in the World List and, for new species, in Table I1 of this review (p. 14). The information is often incomplete because the entire bathymetric range of very few species has been investigated. Mysids, unlike euphausiids, occur in small numbers in oceanic samples but can often occur in vast numbers in neritic and littoral samples. Vinogradov (1970a, b), Vinogradov and Parin (1973), Vinogradov et al. (1970) and Murano et al. (1976) have examined the vertical distribution of the oceanic biomass of mysids and euphausiids. That of mysids tends to be reciprocal to that of euphausiids; mysids are dominant in the littoral and shallow shelf environments, euphausiids are dominant in the epipelagic and upper mesopelagic

4.

VERTICAL DISTRIBUTION AND MIGRATION

69

regions, while mysids are again dominant in the lower mesopelagic and bathypelagic environmenbs. The number of papers defining the vertical distributions of neritic and oceanic mysids are few: Colosi (1930), Taniguchi (1969), Wigley and Burns (1971), Nikolaev et al. (1973), Childress and Nygaard (1974), Murano (1975, 1976, 1977a). Other papers are referred to later in the discussion of the die1 vertical migrations performed by some species. As mentioned above, mysids occur in small numbers in oceanic samples and there is little evidence to suggest that they are important constituents of the deep sonic scattering layers. Populations of shallow water species, however, do occur a t densities such that they can constitute scattering layers. McNaught (1968) discusses the possibilities of observing sonic scattering by populations of the freshwater Mysis relicta in the Great Lakes of North America. Friedl (1971a) concluded that Acanthomysis macropsis and Neomysis kadiakensis appeared to contribute to the records of sound scattering at 38.2 kHz in Puget Sound. Mysids occur to depths of a t least 7210m where Belyaev (1966) recorded Amblyops magna. Bacescu (1971a) found Mysimenzies hadalis at a depth of 6200 m in the Peru Trench and Amblyops aequispina was taken at 5760 m by Birstein and Tchindonova (1958). Bacescu has suggested that further sampling of the hadal region will discover more species of mysids to be present. Wigley and Burns (1971) have presented a bathymetric classification of mysids based upon their studies of the fauna off the eastern coasts of the United States, extending from the littoral zone to the continental slope at depths of 700m. The classification given in Table VII incorporates their findings but modifies the ranges of depths ascribed to their regions and introduces regions outside the scope of their investigations. The first three groups of species, the freshwater, brackish water and littoral are relatively easy to define and recognize in the environment. Shallow shelf species are numerous in number and appear t o form a natural bathymetric group; they are predominantly hyperbenthic in habit. Eurybathic shelf species are probably fewer in number but again form a natural group, and are also predominantly hyperbenthic in habit. Members of the remaining groups are more difficult to recognize in samples from the environment. The deep shelf and upper slope species and the slope species are principally hyperbenthic in habit although the mesopelagic regime impinges on the slope and species can be mis-classified. The catching of a mysid in pelagic as opposed to a hyperbenthic sampler does not necessady define its normal habitat. There are pelagic species occurring over

70

THE BIOLOGY O F M Y S I D S

the continental shelf but they are considered to be relatively few in number and are grouped with the more strictly oceanic epipelagic species. The meso- and bathypelagic species are self-explanatory except in the region of the continental slopes as mentioned above. There is also a possibility that some of the rare deep sea species, occasionally caught in pelagic nets, may be representative of deep hyperbenthic populations. TABLEVII. BATHYMETRIC CLASSIFICATIONOF SPECIES OF MYSIDS

Title of species

Normal habitat

Total range of depth ( m ) ~~

Freshwater Brackish water Littoral Shallow shelf Eurybenthic shelf Deep shelf and upper slope Slope 8. Epipelagic 9. Mesopelagic 10. Bathypelagic 1. 2. 3. 4. 5. 6. 7.

0-50 m depth 0-10 m depth Intertidal or subtidal Less than 50 m depth Broad range of depth Outer shelf and on slope On the slope Oceanic surface layer Oceanic, 200-700 m depth Oceanic, c. 500 m to very deep (6000 m)

~~

0-200 0-20 0-10 2-100 2-400 100-400 200-700 0-300

As already stated, not enough information is available to classify all known mysids into one or other of the classes shown in Table VII. They can, however, be divided between several broad regions (Fig. 19). Region A, brackish, caves and freshwater, contains species in classes 1 and 2 in Table VII. Region B and class 3 are synonymous. Region C includes all shallow shelf, eurybathic shelf, deep shelf and upper slope species ; it probably also includes a few species t,hat should be classed as epipelagic. Region D and class 8 are synonymous. Region E and classes 9 and 10 are synonymous but region E also contains the slope species of class 7. The hyperbenthic habit of many species is reflected by the distribution of stippling in Fig. 19. Mauchline (1972) suggested that deeper living species of mysids tended to be larger in adult body size than shallower living species. This thesis is illustrated in Fig. 20 where the frequency distributions of the maximum body lengths of the various species inhabiting the regions illustrated in Fig. 19 are shown. Data on either body length or depth of occurrence of 24 species are not available and these species are therefore not included in Fig. 19. The majority of species of less

4.

VERTICAL DISTRIBUTION AND MIGRATION

. . ' E

,

.

. .. .

. . ..

71

.

A. Brackish,caves,freshwater

C. Coastal .neritic

D. Epipelagic

E. Meso- and bathypelagic

FIQ. 19. The distribution, shown by the degree of stippling, of the species of mysids in the different regions of the marine environment.

than 5 mm total length occur in the littoral and neritic regions and few species in these regions are greater than 25mm in length. No species less than 5 mm in length is recorded from the meso- and bathypelagic region (including the slope) and an appreciable proportion of these species are more than 25 mm in length; the largest known mysids occur in this region. Many species perform a die1 vertical migration, rising towards or t o the surface layers during the night and returning to the deeper layers at daylight and remaining there throughout the day. The amplitude and form of the vertical migration is limited in the case of littoral and other shallow living species by the depth of water available to them. Mysids generally avoid bright illumination although Praunus $extiasus, for example, can be observed resting on the sand surface facing into the water current in bright sunlight a t a depth of less than 0.5 m; the related species P . neglectus is found resting in these conditions among macroalgae. Macquart-Moulin (1972, 1977a) demonstrated experimentally that Anchialina agilis, Qastrosaccus amatus, Q. lobatus and Siriella jaltensis avoid bright light but may be attracted to weak light. He further showed (1973b, 1977c) that some of these littoral

72

THE BIOLOQY OF YYSIDS

A . Brackish. caves, freshwater

n = 32

lo

----

n = 164

---E. Meso-and bathypelogic

n

=234

45 55 65 75 >80 Body length ( m m ) FIG. 20. The frequency of species of differing maximum body length in the various regions of the marine environment. 5

15

25

35

and shelf species have an endogenous rhythm of swimming activity. Anchialina agilis, frequently living pelagically, showed no clear rhythm and appeared to swim more or less continuously throughout the 24 hours. The hyperbenthic shelf species, Leptomysis gracilis and Siriella jaltensis also showed no clear die1 rhythm of activity. The littoral species S . clausii tended to be more active at night than during the day. The four species, Gastrosaccus armatus, G . lobatus, G. mediterraneus

4.

VERTICAL DISTRIBUTION AND MIGRATION

73

and G. spinifer burrow into the sand during the day and emerge a t night into the pelagic environment; these exhibited the most pronounced diel rhythm of activity. A sexual difference of behaviour was observed in Anchialina agilis where no diel rhythm was present in females but a pronounced rhythm recorded from males. The only deep living species investigated, the upper slope Lophogaster typicus, rested continuously half-submerged in the sediment and showed no Eign of a rhythm of activity. Macquart-Moulin’s experimental observations confirm the conclusions of other investigators sampling wild populations. The following species investigated by the following authors live on or in close association with the surface of the sediment during the day but migrate to or towards the water surface a t night, a distance of up to 50 m : Acanthomysis sculpta by Hobson and Chess (1976) ; Anchialina agilis by Russell (1925), Macquart-Moulin (1965), Champelbert and Macquart-Moulin (1970) ; A. typica by Wigley and Burns (1971) ; Gastrosaccus lobatus by Macquart-Moulin ( 1965), Champelbert and MacquartMoulin ( 1 970) ; G. normani by Russell (1925)) Macquart-Moulin (1965) ; G. psammodytes by McLachlan et al. (1979); G. spinifer by Arntz (1971), Hesthagen and Gjermundsen (1979)) Moller (1979) ; Heteromysis formosa by 0. S. Tattersall (1967); Leptomysis gracilis by Russell (1925), Macquart-Moulin (1965), Champelbert and Macquart-Moulin (1970) ; L. lingvura by Colman and Segrove (1955) ; Metamysidopsis elongata by Clutter (1969); Mysis mixta by Wigley and Burns (1971); M . relicta by Mundie (1959), Beeton (1960), Wells (1960), McNaught and Hasler (1966), Lasenby and Langford (1972), Teraguchi et al. (1972, 1975), Wilson and Roff (1973), Grossnickle and Morgan (1979) ; Neomysis americana by Whiteley (1948), Hulburt (1957), Herman (1963), Hopkins (1965), Wigley and Burns (197 1); N . mercedis (awatschensis) by Heubach (1969) ; Paramysis intermedia and P. ullskyi by Boroditch and Havlena (1973) ; Schistomysis spiritus by Colman and Segrove (1955) ; Siriella chierchiae by Brattegard (1970a) ; S. clausii by Champelbert and Macquart-Moulin (1970) ; S. jaltensis by Macquart-Moulin (1965) ; S. norvegica by Champelbert and Macquart-Moulin (1970) ; S. pacijica by Hobson and Chess (1976). These migrations are probably not performed by the whole population. The investigations are not usually detailed enough to demonstrate this but where several depth horizons were sampled the mysids occurred at all depths during the night. This is a common form of migration in shelf and littoral species, part of the population rising all the way to the surface but a significant portion remaining a t depth. The exact proportion remaining in the hyperben thic habitat is extremely difficult to determine because it

74

THE BIOLOGY OF MYSIDS

involves comparison of the sampling efficiencies of pelagic nets and hyper benthic sledges. Some of the investigations quoted above, for example on Neowysis americana, were made in water columns of less Qhan 15 m depth. These shallow living species are often so closely associated with the sediment during the day that a diel vertical migration can be clearly demonstrated a t night. It can be interpreted as a spreading of the vertical range of the species rather than a vertical migration in which there is an appreciable vertical change in the depth of modal occurrence of the species. There is no evidence suggesting that some individuals migrate to the surface a t sunset and remain there until sunrise while others perform no migration at all. There may be a continuous interchange of individuals, upwards and downwards, within the vertical structure of the population. This would give the impression of a vertical migration when a time series of net samples was analysed but is really an extension of the vertical range of foraging of the population as a whole. A few species do not seem to perform a diel vertical migration and future work may show that many of these species do not migrate. Mysis mixta and Mesopodopsis slabberi are two examples according t o Hesthagen (1973). Mauchline (unpublished) obtained no evidence of a diel migration in Mysidopsis gibbosa or Erythrops elegans in the Clyde Sea area nor in Schistomysis ornata in the Clyde or Loch Etive. Mysidopsis didelphys migrated to a small extent but on many occasions appeared to remain hyperbenthic a t night. Not all populations of a species need necessarily perform a vertical migration. Whiteley (1948) found no evidence of a regular diel migration in Neomysis americana on Georges Bank where the depth at which they were living, as deep as 75 m, was greater than in the coastal regions where this species is known to migrate fairly regularly. The sexual difference in the rhythms of activity of male and female Anchialina agilis noted by Macquart-Moulin ( 1973b) has been observed in the environment. Tattersall (1951) found that male A . typica outnumbered females in samples taken a t night and Brattegard (1970a) made a similar observation and suggested that migration may be associated with sexual activity. Zatkutskiy (1970) and Herman (1963) examining Gastrosaccus sanctus and Neomysis americana found that females carrying embryos or larvae appeared to migrate more regularly than other components of the population. Bacescu (1975), on the other hand, noted that female Gastrosaccus msangii carrying young remained buried in the sand. No consistent differences between the behaviour of male, female and juvenile Mysis relicta were observed by Beeton (1960) although Grossnickle and Morgan (1979) found that

4.

75

VERTICAL DISTRIBUTION AND MIGfCATION

the largest males and females in a population in Lake Michigan occurred deeper than smaller individuals. Studies of the vertical migrations of the meso- and bathypelagic species are relatively few. Waterman et al. (1939) found that Boreomysis microps and Eucopia unguiculata performed a die1 migration (Table VIII) near Bermuda but Pearcy et al. (1977) could demonstrate no such migration in populations of the latter species in the eastern Pacific off Oregon. Pequegnat (1965) found evidence of a migration in the shallowest living Gnathophausia species, G . ingens, that he investigated while Murano (1977a) concluded that a similar migration is TABLEVIII. THEVERTICALMIGRATION OF OCEANIC MYS~DS Average day depth (m)

Average night depth (m)

Total vertical range (m)

Arachnomysis megalops Boreomysis californica B . microps

200-500

200-500

200-500

500-900

500-900

800

B . rostrata

600

Species

Diatance migrated (m)

Reference

0

Murano (19774

0-1000

0

400

200-1200

400

600

0-1000

0

100-700 200-700 0-300 0-500 200-700

100-3000 200-700 0-300 0-500 100-700

P 0

Pearcy et aE. (1977) Waterman et al. (1939) Pearcy et al. (1977) Murano (1977a) Murano (1977a) Murano (1977a) Murano (1977a) Murano (1977a)

400

200-1200

400

700

700

100-1 000

0

Gnathophausia gigas 2100

2100

600-4000

0

G . gracilis

1600

1600

700-3500

0

G . ingens

1100

500

250-2500

400

500-700 500

200-1200 0-2000

400

400-1500

0

Caesaromysis h,iapida 300-3000 Euchaetomera tenuis 200-500 E . typica 0-300 E. glyphidophthalmica 9 Euchaetomeropsis 100-300 merolepis Eucopia unguiculata 800

Katerythrops oceanae 500-700 Meterythrops 900 microphthalma Teraterythrops robUSta -

-

0 0 0

0

Waterman et al. (1939) Pearcy et al. (1977) Pequegnat (1965) Pequegnat (1965) Pequegnat (1965) Murano (1976) Murano (1977a) Murano (1975)

76

THE BIOLOGY OF MYSIDS

performed by Meterythrops microphthalma. Boreomysis curtirostris, B. incisa, B. plebeja and B. semicoeca performed vertical migrations through 4000-5000 m according to Tchindonova (1 959) who found that they had fed on diatoms and surface living crustaceans. Such extensive vertical migrations may not be regular in occurrence but only be performed irregularly. Nemoto (1959) recorded a juvenile Gnathophausia gigas among the stomach contents of a surface feeding fin whale, while Holthuis and Sivertsen (1967) found G. ingem, G . zoea and Chalaraspidum alatum among the stomach contents of an albatross. These latter authors suggest that upwelling of deep water may have been responsible for the presence of these species at the surface and, without further information, this seems plausible. It would appear, therefore, that the majority of deep living species do not perform a diurnal vertical migration to any marked extent. The cost to an animal of a die1 vertical migration in terms of energy expended may be less than previously considered (Vlymen, 1970; Klyashtorin and Yarzhombeck, 1973). Barker (1973) and Foulds and Roff (1976), examining energy expended by Mysis relicta, found no evidence of extra energy expenditure. Teraguchi et al. (1972), however, working with male M . relicta found that they lost about 8% of the calorific content between the start of the evening ascent and the end of the morning descent. Factors that modify the vertical distribution and migrations of mysids are not well documented but they are undoubtedly essentially the same as those that influence the behaviour of other crustaceans. Light intensity appears to be the dominant factor controlling the vertical distribution of mysids (Beeton, 1960 ; McNaught and Hasler, 1966; Heubach, 1969; Wilson and Roff, 1973; Teraguchi et al., 1975). The presence of moonlight or fog influences the extent of the migration. Mysis relicta appears to have increased its depth of occurrence in response to increased penetration of light following the diversion of sewage effluents in Lake Washington (Eggers et al., 1978). The vertical distribution of temperature can have a severe modifying effect upon migration. A sharp thermocline with a gradient of 2'C is an effective barrier to the upward migration of Mysis relicta; migration takes place through weaker thermoclines (Beeton, 1960 ; McNaught and Hasler, 1966). Young mysids were found to be tolerant to vertical stratification of the water, under experimental conditions (Harder, 1968). The vertical distribution of M . relicta in Lake PaajBrvi, Finland is controlled by the position of the 7°C isotherm (Hakala, 1978). Boroditch and Havlena (1973) found that Paramysis intermedia and P. ullskyi, two species that have been introduced t o the freshwater

4.

VERTICAL DISTRIBUTION AND MIGRATION

77

reservoirs of the Volga River, avoid wave action in the litt.ora1 region by moving temporarily offshore to a depth of 2 m. This is true of other littoral species such as Schistomysis spiritus and S. kervillei which move to depths of 2-3 m in exposed situations on the Scottish coast. Flood and ebb tides may also affect the vertical distributions and migrations of some species. Heubach (1969) found that Neomysis mercedis (awatschensis) in the Sacramento-San Joaquin river estuary occurred higher in the water column during both the day and night when the tide was flooding; the flood tides had lower velocities than the ebb tides. Finally, Wilson and Roff (1973) found that My& relicta appears to migrate higher in the water column when the values of surface chlorophyll are high in Lake Ontario. Ontogenetic migrations have been investigated in few deep living species. Casanova (1970, 1977) found that younger Eucopia unguiculata ( E . hanseni) live higher in the water column than older ones. Page (1 941, 1942) presents evidence of ontogenetic migrations in Ceratolepis hamata, Gnathophuusia zoea and Lophogaster spinosus. A similar situation exists in populations of the freshwafier Mysis relicta ; Lasenby and Langford (1972) and Reynolds and De Graeve (1972) found that in vertical water columns of 30 and 80 m respectively adult M . relicta tend to have a modal depth horizon deeper than that of the juveniles. Almeida Prado (1972, 1973) noted that young Bowmaniella brasiliensis and Mysidopsis tortonesei migrate to the surface while the adults remain a t 8 m depth close to the surface of the sediment. A horizontal migration is combined with an ontogenetic vertical migration in a population of Schistomysis ornata studied within a sea loch in Scotland (Mauchline, 1970a). This is illustrated in Fig. 21. The young are concentrated in the innermost region of the loch at depths of approximately 35 m while the mature adults are concentrated towards the seaward end of the loch at depths of approximately 65 m. The region between, which has depths of over 100m present, is sparsely populated by immature juveniles. Hulburt (1957) found a gradient of body size in samples of Neomysis americana over a 20 km length of the Delaware River Estuary; larger individuals tended to occur higher up the estuary than small individuals. Seasonal changes in the vertical distribution of several species have been found. Changes to avoid seasonal extremes of temperature and salinity can be expected in littoral and brackish environments. Kinne (1955) found that, Neomysis integer living in a canal migrated horizontally at first to warmer regions in the autumn. They then moved to deeper but warmer hyperbenthic regions when the warmer shallower areas cooled in the early winter. Mesopodopsis slabberi

-.N Depth (m) 35

lemp. Sol.

("C)

(%o)

10-5 25.0

2km

Depth (m) 110

Body length ( mm ) juveniles;

0immature d a n d ?;

my?with empty marsupium;

* Temp.

Sol.

("C)

(%o)

8.8

=

27.0

??with young;

mature males

4.

VERTICAL DISTRIBUTION A N D MIGRATION

79

appears to exhibit the same behaviour in the Black and Azov Seas (Zatkutskiy, 1970). It is normally regarded as a shallow brackish wa,ter species (Tattersall and Tattersall, 1951) but has a relatively wide bathymetric range of 0-20m during the spring and summer. According to Zatkutskiy, it migrates from the shallow lagoons of the Black and Azov Seas to offshore regions with depths as great as 60 m where it passes the winter. Hesthagen (1973) suggests that Gmtrosaccus spinifer behaves similarly in the western Baltic. It is probable that some littoral and brackish water species will migrate during summer to avoid adversely high environmental temperatures. Many coastal situations subject to freshwater run-off from the land have seasonal periods of low salinity regimes. Praunus jlexuosus, occurring in the littoral region of Loch Etive (Fig. 21), inhabits the inner end of the loch but moves seaward after periods of rainfall have reduced the salinity markedly below 20%,. Teraguchi et al. (1975) examined the changing seasonal response of the freshwater Mysis relicta to illumination. Lasenby and Langford (1972), whom the previous authors do not refer to, found that this species progressively commences its vertical ascent earlier and its descent later during the period spanning the early summer and late autumn. Teraguchi et al. suggest that it has a circannual rhythm in its response to changes in environmental illumination and this modifies the resultant pattern of vertical migration. Pearcy et al. (1977) found that the layer of maximum abundance of tihe bathypelagic Boreomysis californica, within the 500-1000 m depth horizon, occurs about 100 m shallower in winter than in summer. This could be an effect of the changing seasonal altitude of the sun off Oregon resulting in a seasonally changing intensity of ambient light. Mysids live in a wide range of habitats and exhibit a wide range of behaviour. Many species, living in the immediate environs of marine laboratories, are excellent experimental organisms for a variety of purposes. The vertical distribution and migratory behaviour are aspects of the biology of many species bhat require further investigation. Ontogenetic differences in the behaviour of individuals within a population, often reflected in seasonal differences in the behaviour of the population as a whole, compounded with horizontal migrations

FIG.21 (opposite). Population histograms of samples of Schistomysis ornata taken from different regions of Loch Etive, Scotland, in November, 1967. The positions of the samples are shown by the bars; the 25, 50 and 100 m depth contours are indicated by the dotted lines. Depth, salinity and temperature of the water where the samples were taken are shown.

80

THE BIOLOGY OF MYSIDS

of aggregations or swarms that cause difficulties in location of the animals a t subsequent sampling dates, make studies of their vertical distribution difficult. Nevertheless, studies of the behaviour of these animals would aid considerably in an understanding of that of less accessible organisms.

CHAPTER 5

THE GUT, FOOD AND FEEDING Mysids live in a wide range of habitats with a diversity of food resources that they exploit very successfully. This is reflected in the different forms of the feeding appendages. These have been modified in a number of ways throughout the Order but the functional significance of many of the modifications is unknown. The feeding appendages of Gnathophausia xoea, Lophogaster typicus and Hemimysis lamornae have been studied in detail by Manton (1928s) and Cannon and Manton (1927a), who review the earlier work. The general arrangement of the appendages is described in mysids by these authors and in euphausiids, in which they are analogous, by Mauchline and Fisher (1969). They consist, in anterior to posterior order, of an unpaired labrum and paired mandibles, paragnaths (labia), maxillules and maxillae. The thoracic legs also frequently function in feeding. The labrum is single, anterior to the mouth, and projects ventrally. The biting surfaces of the mandibles fit against its posterior surface. The paragnaths are relatively simple in form, plate-like and project ventrally, fitting closely against the posterier faces of the mandibles. Next are the maxillules and maxillae followed by the thoracic legs. The labrum and paragnaths form a chamber ventral to the mouth and cutting surfaces of the mandibles, which constitute the lateral walls of this chamber and macerate food within it. The appendages of the different species of mysids have been illustrated many times in the taxonomic literature. No comprehensive, comparative study however, of their morphology relative to their function has been made. The mandible has cutting, grinding and macerating regions and also a forwardly directed mandibular palp (Fig. 22). The general form and proportions of the mandible and its palp can vary considerably. Those of Schistomysis ornata are of a general form found throughout the Order while that of Petalophthalmus armiger is an extreme form with a much enlarged and modified palp. The terminal segment of the palp often has a cleaning mechanism of the form found in Schistomysis ornata (Fig. 22); this is presumably used to clean the cutting

82

THE BIOLOOY OF MYSIDS

FIQ.22. Right mandible of (A) Schistomysis ornatu and (B) Petalophthalmw armiger. cm, cleaning mechanism.

and grinding surfaces of the mandible as well as to remove food from the more posterior mouthparts and push it into the mouth. The morphology of the cutting and grinding surfaces varies between species quite considerably (Fig. 23). The left and right mandibles are asymmetrical, as illustrated in Neomysis rayii (Fig. 23) and Gordon (1964) points out that, in consequence, both mandibles should be drawn. This asymmetrical design allows the cutting surfaces to dovetail into one another. All mandibles illustrated in Fig. 23, with the exception of those of N . ra ii, are those of the right-hand side viewed from the anterior aspect. aking that of Boreomysis microps as an example, the cutting

4

' I

F D

H

G

J FIG.23. Cutting edges of the right mandibles of A, Gnathophawk zoea; B, Lophogaster typicus; C , Eucopia sculpticauda; D, Boreomysis microps; E, Meterythrops p i c k ; F, Antarctomysis maximu; G , Praunua $ e m o m ; H, Longithorax f w c w ; J, Neomysis rayii showing asymmetry of right and left mandibles. c, cusp, synonymous with lacinia mobilis; ip, incisor process;'lm, lacinia mobilis; pm, pars m l a r i a ; 81, spine row.

84

THE BIOLOGY OF MYSIDS

edge includes a dorsal grinding region, the pars molaris, very large in this species. This region is' adjacent to the mouth. An extensive row of prominent spines, the spine row, is present in the lower and central region. Ventral to it is the lacinia mobilis, spines which are movable, and the incisor process. Not all of these elements are present in other species. The left-hand mandibles of Gnathophausia, Lophogaster and Eucopia species have a lacinia mobilis but the right-hand mandibles only have a fixed cusp (Fig. 23). The spine row is simple in the above genera of lophogastrid mysids and almost completely absent in Longithorax fuscus, but highly developed in Meterythrops picta. The grinding area of the pars molaris is reduced in size in Lophogaster typicus and absent in Meterythrops picta and Longithorax fuscus. It is also reduced in size or absent in species of Amuthimysis, Arachnomysis, Echinomysides, Mysidopsis, Heteromysis and Stygiomysis. The commonest forms of mandibles found throughout the Order are similar to those exemplified by Antarctomysis maxima, Praunus jlexuosus and Neomysis rayii (Fig. 23). The size of the pars molaris relative to the rest of the mandible is related to the diet of euphausiids (Mauchline, 1980). A large pars molaris is associated with herbivorous feeding, the hard diatom frustules being ground by this region. This relationship is probably true in many species of mysids, in, for example, the genera Praunus, Antarctomysis, Neomysis and Acanthomysis. The function of a large grinding region in mandibles of deep living species where a diet of diatoms is not obviously available may be different. Tchindonova ( 1 959) examined and figures the mandibles of Gnathophausia gigas, Eucopia australis, Boreomysis plebeja, B. sibogae, Meterythrops microphthalma, Dactylamblyops laticauda and Petalophthalmus armiger. She examined the stomach contents of a wide range of oceanic species and found that the stomachs of Boreomysis curtirostris, B. incisa, B. plebeja and B. semicoeca caught in the 2000-4000 and 4000-7500 m horizons contained slightly digested remains of diatoms and surface living crustaceans that they had eaten during their vertical migration. Consequently, the presence of a well developed pars molaris in a supposedly deep living species may indicate that it obtains phytoplankton in significant quantities either directly or indirectly from the surface layers. The labium or lower lip of mysids is cleft, the ventrally projecting plates being known as a paragnaths (Fig. 24). These lie closely against the posterior faces of the mandibles. Gordon (1960) examined the paragnaths of a variety of species because those of Stygiomysis holthuisi (Fig. 24) are peculiar in that they are spoon-shaped rather than

5.

THE CUT, FOUL) AND FEEDINC:

B

85

B

E

FIG. 24. The paragnaths of the labium of A, Neomysis ‘czerniavskii;B , Gnatlwphausio zoea, anterior view showing asymmetry of paragnaths; C, ct. zoea, posterior view showing how left paragnath fits into the recess in the incisor region of the mandible; D, Pekclophthalmus arrniger ; E , Stygiomysis holthuisi ; F, Boreomysis Tostrata (Figs D-F after Gordon, 1960).

rectangular like the more typical paragnaths of Neomysis czerniavskii (Fig. 24). Clarke (1961a) describes the asymmetrical paragnaths of Gnathophausia species and points out that such paragnaths also occur in at least some species in the genera Chalaraspidum, Pseudochalaraspidurn, Lophogaster, Paralophogaster, and Eucopia. Those of Gnathophuusia zoea are shown in Fig. 24. The left paragnath is more developed and is armed with tubercles and ridges. It fits into the posterior face of the left mandible which is curved in the incisor region to receive it (Fig. 24). The right paragnath is thin, conventional in form and is not coupled with the right mandible. The paragnaths have no muscular system and the mandible induces movement in the left paragnath whose tubercles and ridges then form an integral part of the mandible.

86

THE BIOLOGY OF MYSIDS

The functional significance of variations in the shapes of the symmetrical paragnaths of the mysids is unknown. The form of the maxillule in a variety of species is shown in Fig. 25. It is relatively simple, consisting of a proximal and distal endite. The distal endite invariably has a row of more or less strong spines while the proximal endite is armed with setae and three or four long spines. The posterior face of the distal endite often has a group of three or more strong setae, shown hatched in Fig. 25. The appendage

A

D

C

B

E

F

FIG. 25. The right maxitlule of A, Eucopia sczclpticauda; B, Meterythrops picta; C , Longithorux fuscus ; D, Antarctomysk maxima ; E, Schistomysis ornata; F, Praunus Jexuosus. de, distal endite; pe, proximal endite.

is the most constant in form and, therefore, probably in function. One variation that exists in relatively few genera is a reduction in the number of long spines, as opposed to setae on the proximal endite. Species in the genera Mysidopsis, Metamysidopsis, Promysis and probably some species of Lophogaster and Siriella have only two spines while Pillai (1973) figures this endite of Siriella jonesi with one spine. The only other marked variation in form occurs in Mysidella species where the spines on the distal endite decrease in size dorsally round its curved edge. The maxilla can vary considerably in form, especially in the relative proportions of the exopod and terminal endopodite and in the setulation of the appendage as a whole (Fig. 26). The maxilla of Lophogaster typicus has long weak setae and no spines; the form of maxilla

1

I

e

C

'

\\\

FIQ.26. Right maxilla of A, Lophogaster typicus; B, Meterythrops picta; C, Boreomys~s microps; D, Longithorax fuscus; E, Eucopia sculpticauda ; F, Petalophthalmua armiger; G , Praunus fiexuosus; H, Antarctomysis nutxima. e, exopod; 8 , spine on proximal lobe.

88

THE BIOLOGY OF MYSIDS

found in this species is uncommon among mysids. The maxilla of Meterythrops picta is of a form typical of other species in the tribe Erythropini; it is heavily setose with a few spines present. The maxilla of Boreomysis microps has many fine filtering setae and a row of spines is present on the internal (lateral) edge of the terminal endopodite; such a maxilla occurs in many species of the tribe Erythropini and also in species in the genera Hansenomysis, Siriella, Dioptromysis and Heteromysis. The type of maxilla found in Longithorax fuscus also occurs in some species of Siriella, Anchialina and Gastrosaccus; it is lightly armed with plumose setae and has few, if any, spines. The maxilla of Eucopia sculpticauda is closely similar to those of species in the genera Gnathophawia, Anchialina and Mesopodopsis ; the terminal endopodite is enlarged and the appendage is heavily armed with plumose setae but with few spines. The form of maxilla found in Yetalophthulmus armiger is rare and may be restricted to this genus; it is fragile and armed with weak setae and a long terminal spine on the terminal endopodite. The maxilla of Praunus Jlexuosus is typical of those of many species in the tribe Mysini but also occurs in species in the genera Dactylamblyops, Eoerythrops, Pseudomma, Leptomysis, Mysidetes and Spelaeomysis; it is characterized by a long strong spine on the proximal lobe (Fig. 26) and a row of spines on the inner (lateral) edges of the Oerminal endopodite and the other lobes. The terminal endopodite of the maxilla of Antarctomysis maxima has a more extensive and ventrally situated row of spines which is present in this region of the maxillae of species in the genera Bathymysis, Mysis and Hemimysis. This row of spines is even more accentuated in the maxillae of species in the genera Afromysis, Doxomysis, Hyperiimysis, Iimysis, Nouvelia, and Pseudoxomysis (see Figs 7.27, 7.29-31, 8.3). The only major form of maxilla not illustrated in Fig. 26 is a maxilla of the general form of that of Longithorax fuscus but with relatively few short setae interspersed between relatively many sharp spines, arranged and directed internally towards those of the opposite appendage. This maxilla occurs in species in the genera Cubanomysis, Mysidopsis, Metamysidopsis and Promysis of the tribe Leptomysini and also in species of Stygiomysis and Spelaeomysis. The form of the thoracic appendages varies and their morphology is documented in detail in the taxonomic literature. They have a welldeveloped endopod and exopod. The exopods itre of the general form shown in Fig. 27A, B and may rarely, in the first thoracic leg, be reduced as in Longithorax fuscus (Fig. 27C), or absent. The form of the endopods is more variable. That of the first thoracic leg is usually relatively shorter than those of more posterior legs. It may be of the

5.

8!,

T H E UUT, FOOD AND FEEDING

relatively simple form found in L. fuscws or it may have well-developed lobes on the second, third and fourth segments as in Schistomysis ornata (Fig. 27). The endopods of more posterior thoracic legs are usually of the form illustrated for the second leg of S. ornata. The morphology of the terminal segments of the endopods can vary. The terminal segment of either one or both of the first and second legs may have a cleaning mechanism similar in structure to that on the

en

e

E Fro. 27. Thoracic legs. A, first thoracic leg and B, second thoracic leg of Schistomysia ornata; C, first thoracic leg of Longithorarfwcus; D, terminal segment of the endopod of the third thoracic leg of Eucopia sculpticauda; E , terminal segments of the endopod of the third thoracic leg of Heteromysis nzicrops; F, enlargement of one of the structures indicated in the penultimate segments of the leg of H. microps (E and F after Tattersall and Tattersall, 1951). en, endopod, ex, exopod, 1, lobe.

distal segment of the mandibular palp of S. ornata (Fig. 22, p. 82). These segments may terminate in a nail (spine) or a series of spines as instanced in Longithorax fuscus and Eucopia sculpticauda (Fig. 27), and in Petalophthalmus armiger. The presence of a terminal nail or spine, usually of a less robust form than that of Eucopia sculpticauda, on the terminal segment of the endopods of the thoracic legs is a

00

THE BlOLOCY OF MYSIDS

general feature of mysids. It occurs on the endopods of a t least the third to eighth thoracic legs of most Lophogastrid species, of species of Boreomysis, Siriella, the tribes Erythropini and Leptomysini, some genera of the tribe Mysini, and Heteromysini. They are much less developed in genera such as Gastrosaccus and Anchialina, Parampsis and Schistomysis, and Mysidella. In genera such as Eucopia and Hansenomysis, the terminal nail and one or more setae on some endopods form a pseudochela. Several species of mysids have one or more pairs of the thoracic endopods especially modified. One of the most spectacular modifications is the enlargement of third endopod of Heteromysis microps (Fig. 27) which is larger than the whole cephalothorax. The penultimate segment, the merus, has ten peculiar structures with a barbed seta emerging from a terminal socket (Fig. 27). The nail and two terminal projections (spines) form a pseudochela. Elongation of one or more pairs of thoracic endopods is not a common feature of mysids. The fifth to eighth pairs are elongated relative to the others in Eucopia species while the third pair alone is elongated in Wemisiriella species. The funct;ional aspects of these modifications of the mouthparts and thoracic appendages of mysids are, for the most part, unknown. The only detailed study of how the appendages are used in feeding is that of Cannon and Manton (1927a) on Hemimysis lamornae and some earlier studies referred to by them. This species might be described as a conventional or generalized mysid whose appendages are of a form common throughout the order. It feeds, according to Cannon and Manton, by several methods. One way is by a modification of the generalized filter feeding method that is presumably used by the animal when it is not in direct contact with the surface of the sediment. The modification involves the animal standing on its head on the surface of the sediment and lifting the surface material into suspension by creating a water current with its thoracic exopods. This material is then filtered from the water by the appendages. Another method is used to feed on large particles, a method often described as raptorial. Dead carcasses can be lifted from the surface of the sediment by the thoracic endopods and manipulated against the mouthparts, primarily by the mandibular palps and the endopods of the first two pairs of thoracic legs. The anterior thoracic endopods in conjunction with the mouthparts in many species form a “food basket” analogous with that described in euphausiids (Mauchline and Fisher, 1969). Organic matter from the surface of the sediment is collected by the appendages, especially the mandibular palp, and held in the “basket” prior to maceration by the mouthparts. Gastrosaccus simulans

5.

THE OUT, FOOD AND FEEDING

91

appears to be a strict filter feeder (Nath and Pillai, 1976), feeding on diatoms and dinoflagellates and some algal filaments, although it becomes carnivorous in the laboratory. Mysis relicta is considered by Bowers and Grossnickle (1978) to be a major grazer of phytoplankton in Lake Michigan. Lophogaster typicus according to Manton (1928a) is incapable of filter feeding and the modification of its mouthparts facilitates feeding on large lumps of food material on the surface of the sediment. Molloy (1958) observed Praunus Jtexuosus browsing on the epifauna of seaweeds. Some species of mysids are carnivorous and Lasenby and Langford (1973) observed Mysis relicta attacking and eating live Daphnia. Tchindonova (1959) and Vinogradov (1962) state that Petalophthalmus armiger and Longithorax fuscus catch their prey and suck out the internal contents, a habit known to occur among euphausiids feeding on copepods (Mauchline and Fisher, 1969). Molloy (1958) found that Neomysis integer and Praunus jlexuosus, feeding on dead carcasses of mysids, preferred the thoracic to abdominal region of the carcass. Once the food has been gathered and macerated by the appendages it is passed to the mouth and eaten. Gelderd (1907) describes the histological structure of the alimentary canal of Anchialina agilis, Gastrosaccus spinifer, Mesopodopsis slabberi, Mysidopsis gibbosa, Neomysis integer, Praunus jlexuosus, Schistomysis ornata, S . spiritus and a species of Siriella. Siewing (1953) illustrates the internal armature of the stomach of Eucopia sculpticauda. Molloy (1958) has re-examined some of these species but added others ; she studied Anchialina agilis, Gastrosaccw spinifer, Hemimysis lamornae, Leptomysis mediterranea, Mesopodopsis slabberi, Neomysis integer, Praunus jlexuosus, Siriella armata and 8. jaltensis. More recently, Nath and Pillai (1972, 1976) examined the structure of the canal in Spelaeomysis longipes and Gastrosaccus simulans. The alimentary canal of mysids has the same general structural plan as that of the decapods. The arrangement of the alimentary tract is shown in Fig. 28. The oesophagus is short and muscular, lined with chitin and has spines or setae that tend to point upwards towards tihe stomach. The stomach is divided into an anterior cardiac region and a posterior pyloric region. Both regions are usually armed internally with spines and setae. The detailed form and armature of the stomach varies between species. The stomach of Neomysis integer (Fig. 29) is typical of that of many species. The anterior cardiac region of the stomach of Mesopodopsis slabberi seems to be exceptionally small relative to the pyloric region. The latter species, however, has an unusually long oesophagus. The armature of the stomach of Hemimysis lamornae, Neomysis integer

92

THE BIOLOGY OF MYSIDS

FIQ.28. Generalized diagram of the digestive glands and anterior region of the alimentary canal of a mysid (after Molloy, 1958). cr, cardiac region of stomach; dd, dorsal diverticulum; dg, digestive glands shown stippled; i, intestine; oe, oesophagus; pr, pyloric region of stomach.

and Praunus Jlexuosus is strong but less strong than that found in the stomachs of Leptomysis mediterranea, Siriella armata and S. jaltensis. These armatures contrast strongly with the markedly reduced armature of the stomach of Mesopodopsis slabberi, studied by both Gelderd (1907) and Molloy (1958). Nath and Pillai illustrate a heavy armature of the stomach in Gmtrosaccus simulans but a lighter armature in Spelaeomysis longipes. Mysids have a single or paired dorsal diverticula that arise as extensions of the mid-gut in the posterior dorsal area of the pyloric region of the stomach (Fig. 29). The peritrophic membranes, within which the faecal material is bound, are continuously secreted by them into the intestine. The diverticulum is relatively large, but single, in Hemimysis lamornae, Neomysis integer, Praunus Jlexuosus, Schistomysis ornata and S. spiritus but are smaller and paired in Leptomysis mediterranea, Mysidopsis gibbosa, Mesopodopsis slabberi and Siriella armata (Gelderd, 1907; Molloy, 1958). Both Molloy and Gelderd found that the diverticula are paired in Gmtrosaccus spinifer while Nath and Pillai (1976) found a single diverticulum in G . simulans. These latter authors found that Spelaeomysis longipes also has a single diverticulum and that it is extremely long, reaching the anterior end of the stomach. The digestive gIands consist of two groups of five digitiform processes, one group on each side of the body (Fig. 29). They open by a common duct from the posterior ventral area of the pyloric region of the stomach, the longest pair of elements reaching as far posteriorly as the fourth abdominal segment. The intestine emerges from the posterior end of the pyloric region of the stomach in approximately the third or fourth thoracic segment.

A

Fro. 29. Longitudinal sections of the stomachs of A, Neomysis integer and C, Mesopodopsis slabberi. B, a transverse section of the cardiac region of the stomach of Hem.imysis Zamomae (after Molloy, 1958). cr, cardiac regions of stomach; dd, dorsal divert~uulum;m, mouth; oe, oesophagus; pr, pyloric region of stomach.

94

THE BIOLOGY O F MYSIDS

It extends posteriorly as the mid-gut to the posterior end of the fifth abdominal segment where it becomes the hind-gut lined with chitin. The hind-gut is thicker in diameter and more muscular than the midgut, the detailed histology of the whole intestine being described by Gelderd (1907) and Nath and Pillai (1972, 1976). The mid-gut of the freshwater subterranean species, Spelaeomysis longipes, has posterior caeca that are the site of calcium storage. This calcium is accumulated during premoult stages because it is not readily available in the environment (Nath, 1972). The intestines of mysids exhibit retroperistalsis (antiperistalsis) which is more or less continuous in some species and not in others. Molloy (1958) found that these contractions were present more or less continuously in Praunus Jlexuosus, whether faecal material was presenb in the abdominal region of the intestine or not. Retro-peristalsis in Neomysis integer, on the other hand, seemed to be initiated when faecal material passed into the abdominal region of the intestine. Faecal material, however, was able t o move posteriorly along the intestine even in the absence of any obvious retro-peristaltic contractions (Molloy, 1958). Vigorous retro-peristaltic contractions start when the faeces enter the wider hind-gut region, continue until the faeces are successfully voided, and then cease shortly afterwards. Fox (1952) found that Hemimysis lamornae and Siriella arrnata, in common with many crustaceans, draw water into the gut both through the mouth and anus. The function of such oral and anal intake of water is probably to stretch the gut wall so that the muscles are toned to act rhythmically and effectively. Molloy noted that the food required 30-40 min, occasionally as much as 1.5 h, to pass along the complete length of the alimentary tract. The food was in the stomach and anterior region of the midgut for most of this time. A faecal string, once formed in the thoracic region of the mid-gut, usually passes out of the anus within a few seconds. Part, a t least, of the food passes from the pyloric region of the stomach into the lumina of the digestive glands where it is circulated from one tube to another via the lower pyloric region. The walls of the digestive glands contain circular muscles which power the movement of food and digestive enzymes within the lumina. The digestive glands not only produce digestive enzymes but the finely particulate food is digested and absorbed there. They act as storage tissues for lipids and possibly glycogen; Molloy did not find marked quantities of glycogen but, as Raymont et al. (1968) point out, this is used rapidly and would probably only be detected in freshly caught animals. Molloy found that it was present in the glands when the diet contained excess carbohydrate.

5.

THE GUT, FOOD AND FEEDING

95

The stomach contents of various species of mysids have been examined (Table IX). The food of mysids, like that of many other types of crustaceans, is macerated so that stomach contents primarily consist of unidentifiable, finely particulate matter. The main conclusion that can be drawn from examination of the stomach contents of species is that they eat a great variety of organic materials in their environments. The two Mysidopsis species (Table IX) are rather exceptional i n that their diets would appear to be dominantly carnivorous. On the other hand, Gastrosaccus simubns appears to be a strict filter feeder (Nath and Pillai, 1976) and Mysis relicta can undoubtedly exploit the spring bloom of the diatom Melosira sp. in Lake Michigan although its diet is more catholic a t other times (Bowers and Grossnickle, 1978). Muus (1967) suggests that dense shoals of Neomysis integer in a Danish fjord were feeding on the organic effluent from a slaughter house. Ontogenetic and seasonal changes can be expected in the diet of many species although these have not been investigated, with one exception. Kost and Knight (1975) found that diatoms become progressively less important while detritus becomes progressively more important in the diet of Neomysis mercedis as the youngest juveniles of 2 mm body length grow to adults of 15 mm body length. Brown and Talbot (1972) suggest that Gastrosaccus psammodytes feeds more commonly a t night than during the day. Die1 feeding rhythms are probably present in some species and not in others. Siriella pacijica is quite clearly a nocturnal predator among the kelp beds of California, feeding on copepods and cladocerans, while Acanthomysis sculpta feeds by both day and night on Macrocystis pyrifera and crustaceans (Hobson and Chess, 1976). Assimilation of food by mysids has not been studied in any detail. Blegvad (1922), feeding Praunus species a mixture of mussel flesh and algae, found that 42-113 g dry weight food produced 1 g dry weight mysids. Mysids have been maintained experimentally in the laboratory but on a restricted range of foods. Pechen’-Finenko and Pavlovskaya (1975) fed Neomysis mirabilis on diets of vegetable or animals’ detritus, melanin and dinoflagellates. They fed most voraciously on Gymnodinium kowalewskii and preferred animal to vegetable detritus. Murano ( 1 9 6 6 ~ )found that N . intermedia ate live Artemia and daphnids but would not eat the yolk of chicken eggs or fish meal. They survived better if the bottoms of the culture vessels were covered with mud or sandy mud. Lasenby and Langford (1973) determined the assimilation efficiencies for different foods in Mysis relicta. Daphnia pulex was assimilated with an efficiency of about 85% and chironomid

96

THE BIOLOGY OF MYSIDS

TABLE IX.

Species Acanthomysis aculpta Antarctomyais maxi^ Borwmyais sp. Erythrops elegans Erythrops serrata Gastroaaccua psammodytes Qaatrosaccua apinifer Leptomyais gracilia Mesopodopais slabberi Mysidopais didelphya Mysidopais gibbosa Mysis mixta Myaia relicta Neomyais integer Neomysis intermedia Neomyais mercedis Paramyais arenosa Praunua JEexuosUs Praunua neglectua Praunus spp. Schiatmyaia kervillei Schistomyais ornata Schistomysia spiritua Siriellu paci$ca Spelaeomyaia longipes

DinoDetri&geltus Algae D i a t o m lates

TintinPolynida Rotifers Sponges chaetes

X

x

X X

X

X X

X

X

x

x

X X

x X X

X X

X

X

X

X

x

x

X

X

X

X

X

X

X X

X

x x

x x

X

X X X

X

X

X

X

X

X

X

X X

X

X

X X

X

X

1, Apstein, 1906; 2, Blegvad, 1922; 3, Depdolla, 1923; 4, Lucas, 1936; 5, Tattersall end 1968; 10, Mauchline, 1969a; 11, Mauchline, 1970a; 12, Mauchline, 1970b; 13, Mauchline, 19718; 1972; 18, Neth end Pillai, 1972; 19, Nemoto, 1972; 20, Lesenby and Langford, 1973; 21, Kost and Groesnickle, 1978.

larvae with an'efficiency of 82%. Clutter and Theilacker (1971), using a mysid that they do not name, fed phytoplankton and estimated the assimilation efficiency to be 90%. Mysis stenolepis is able to assimilate sterile raw cellulose with efficiencies of 30-50y0 and sterile hay with efficiencies of 20-35y0 (Foulds and Mann, 1978). The presence of micro-organisms on the cellulose and hay surprisingly decreases the assimilation efficiencies. Foulds and Mann suggest that a gut microflora may be involved in the breakdown of the cellulose. Shushkina (1972) determined K,, the percentage of assimilated food that is converted into new tissue, in Neomysis mirabilis as close to 57%.

5. THE

97

OUT, FOOD AND FEEDINQ

FOODOF MYSIDS ~

Terrigenous Mol- Ostra- Clado- Cope- Amphilwca cods pods pods Crustacea Carrion Material Insects cera X X

Faecal Pellets

X X X X

x

X

X X

X

X X

X X

X

X

X

X

X

X X

X

X X

X

X X

X X

X

X X

X X X

X

X

X

X

x

x

X X

X

X

X

X X X

X

Referencm 22 23 19 9 9 17 2 10 6 12 12 1 5, 20 4, 6, 14 7 21 13 3, 15 15 2 16 11 5, 8 22 18

Tattersall, 1951; 6, Kinne, 1955; 7, Murano, 1963b, 1966c; 8, Mauchline, 1967; 9, Mauchline, 14, Mauchline, 1971b; 15, Mauchline, 1971c; 16, Mauchline, 1971d; 17, Brown and Talbot, and Knight, 1975; 22, Hobson and Chess, 1976; 23, Mauchline, unpublished; 24, Bowers

Pasternak (1977) extended this study and shows that K, changes with. age (body size). It has values of 70-80% .in early juveniles of N . mirabilis but these decrease t o 15-30% in the adults.

CHAPTER 6

CHEMICAL COMPOSITION The chemical composition of mysids has been investigated in recent years but centres on relatively few species. The genus Neomzjsis, which includes many shallow-water, estuarine and so readily accessible species, has received particular attention ; this is especially true of the north-eastern Atlantic species N . integer investigated primarily by Raymont and his collaborators. The results of proximate analysis (Table X ) of mysids are closely similar to analogous data on euphausiids. Dry body weight approximates closely to 20% of the body wet weight with the exception of an adult female Gnathophausia ingens. This is g, bathypelagic species and Childress and Nygaard (1974), in a comparative study of Crustacea, found that shallow living species (0-150 m) and deep living species (900-1200 m) have significantly more water present in their bodies than species living at intermediate depths. This is a broad generalization drawn from moderately variable data. Ash, chitin and the low values for carbohydrate are as expected (Table X) from analyses of comparable organisms. The ash content represents about 1*5-2*5y0 of the wet weight and consequently about 7-5-12.5% of the dry weight. Fukai et al. (1962) determined 8.3% of the body dry weight of Neomysis japonica, Fujika et al. (1969) 5.6% of the dry weight of a Neomysis sp. as ash, and Tracy and Vallentyne (1969) state that the freshwater Mysis relicta contains 21% ash and 15.6% lipid on a dry weight basis. Small sexual differences in body composition were demonstrated in Taphromysis bowmani ; males have significantly more protein and ash and significantly less chitin than females but there are no differences between the carbohydrate and lipid contents (Johnson and Hopkins, 1978). The variation in values for fat and protein are discussed later. Simultaneous seasonal variations in these constituents are described in N . integer by Raymont et al. (1966), in N . awatschensis by Chin (1974), and in Taphromysis bowmani by Johnson and Hopkins (1978); these sets of data are all on a dry weight basis. Elementary analysis of mysids are few and little can be concluded

'TABLE x. PROXIMATE ANALYSESO F

Species Gnathophausia ingens G . ingens (adult 0 ) G . gigas a. gracilia Qnath0pham.a sp. E w p i a sp. Boremy& culijornica Siriella aequiremia Leptomyais lingvura Metamydops-is elongataa Mys-is atenobpis M . relicta Praunua Jexuoaua P . negbctua MesOpodOpd8 slabbe& Neomys-ia integer N.a m h n a N . intermedia N . awatachemk Taphromysis bowmania

MYSIDS

(%

WET WEIGHT)

Water

Ash

Chitin

Carbohydrate

Fat

Protein

72.8 f 0.7 90.7 79.5 f 2.0 81.6f 0.4

3-90 2.65 3.77 3.81

2.54 0.76 2.15 1.18

82-6f 0-9 81.3 70.0

3.31 1.90

0-61

0.14 0.03 0.10 0.13 0.6 0-6 0.1

10.5 2-6 5.4 5.8 4-0 15.0 4.6

8.6 2.3 6.3 5.5 8.0 22.0 4.9

2.50

1.40

0.7-0.9 0.3

2.9-3.5 2.0

20.3-21.6 14.0

79.0 f 3.0 76.3 74.3-7 7.8 88.0 78.3 78.0-85.4 80.2 76-82

1.70-2.2 1'9-3.7

1.6 f 0.2 0.07-0.90

1.6-4.3

1.40

0.1-0.3 0.2-0.5 0.2

2.63 1.40 3-3-4.0

1.40 1.3-1.9

0.3-0.5 0.35-0.50

3.2-5.9

20.7f0.6

2.0 0.2-4.9

13.0-14.0

1.6-2.4 2-7 2-8-3.0

14.9 14.1 11-1-11.6

References 17 17 17 17 10 10 17 9 5 11 15 19,21 8, 18 4 1,4, 6, 7, 13, 18 2 3, 14 12, 16 20

0 Dry weight values transformed to wet weight by assuming dry weight is 20% of wet weight. References: 1, Raymont and Krishnaswamy, 1960; 2, Raymont and Conover, 1961; 3, Hirano et al., 1964a, b ; 4, Raymont et al., 1964; 6,Linford, 1965 and Raymont and Linford, 1966; 6, Raymont et al., 1966; 7, Raymont et al., 1968; 8, Seguin, 1968; 9, Omori, 1969a; 10, Raymont et al., 1969; 11, Clutter and Theilacker, 1971; 12, Lee and Chin, 1971; 13, Srinivasagam et al., 1971; 14, Teshima and Kanazawa, 1971; 15, Tyler, 1973; 16, Chin, 1974; 17, Childress and Nygaard, 1974; 18, Moore, 1976; 19, Salonen et al., 1976; 20, Johnson and Hopkins, 1978; 21, Yaguchi et al., 1974.

100

THE BIOLOGY OF MYSIDS

from them as yet, except that they appear closely similar to those of euphausiida, about wliich more is known. Vinogradova and Petkevich (1967) examined the concentrations of 25 elements in the ashes of Mysis microphthalma and of mixed samples of this species and Paramysis loxolepis (Table X I ) ; their data on these Caspian Sea species have been converted from a yo ash weight basis to an approximated mg% dry weight basis in Table XI by assumjng that the ash TABLEXI. ELEMENTARY COMPOSITION OF MYSIDS (MG/100 0 DRYW E I G H T )

Element

Neomysis integer

Neomysis Mysis Mysis microphthalma intermedia microphthalma Paramysis loxolepis 1140 51

Na Mg

Al Si

K C&

Ti Cr Mn Fe Ni

1050" 1.2b 47.6b

cu

Zn Ga Sr

497 750

26.5 6.1

3"

188 125 113 12.5 313 63 113 2-5 14 25 0.1 125

Ag SZl

Ba Pb

1-3 125 1.1

12-376 62-113 31-150 3-12 187-31 3 2 0.6 3-39 19-25 4-18 12-38 0.1 37-113 1.0

0.3 12-50 0'3-1.2

Data from: Fukai et aE., 1962; Hirano et al., 1964a, b; Raymont et al., 1964, 1966; Vinogradova and Petkevich, 1967; Fujika et al., 1969; Omori, 1969a. a Nemnysis japonica.

bNwmyak sp.

content represents 12.5% of the dry weight. Thommes et al. (1972) measured the mercury content of Mysis reticta in the'Great Lakes of North America in the years 1967-1968 and recorded 0.003 mg Hg/ 100 g wet weight. Mysis relicta, M . mixta, Neomysis integer and Praunus Jexuosus had 0-006-0.008 mg Hg/lOO g wet weight in 1971-1973 in the Baltic Sea (Nuorteva and Hasanen, 1975). The tantalum content of Neomysis integer was 0.23 mg/lOO g dry body weight; other invertebrates had lower concentrations, 0.001-0.024 mg/100 g dry weight

6. CHEMICAL COMPOSITION

101

(Burton and Massie, 1971). Leatherland et al. (1973) found the following concentrations per 100 g dry body weight of the following elements in Eucopia sculpticauda taken from the entrance to the Mediterranean in October, 1970; 3.0 mg arsenic; 0-2 mg cadmium; 0.0011 mg antimony; and 0-02mg mercury. The concentrations of many metals vary considerably within marine organisms in time and space because there are often major contributions of them to the environment from industrial effluents. Analyses of carbon, nitrogen, hydrogen and phosphorus have been made ir, the species shown in Table XII, and are within the expected TABLEXII. CARBON,NITROGEN, HYDROGEN AND PHOSPHORUS IN MYSIDS (yoDRYWEIGHT)

Speciea

C

N ~~

cfnathophazcsia ingens G . ingem (adult) c f . gigas

gracilia Lophogaster sp. Boremy& ealifornka Siriella aequiremia Metmysidop8is elongata whole body cf.

49.2 36.8 41.3 66.2 41.9 46.8 43.4 42.4

cest

36.8 23.5

egg

68.0

larva

Mysis relicta N e m y s i s integer N . internzedia

45.7 50.0 22-35

H

P

~

6.3 6.4 7.1 6.9

8.1 6.1 6.8

7.0

6.9

7.9 11.0

6.5

6.7

11.5

10.9 11.4

2.0-2.5 1.2

Data from: Curl, 1962; Hirano et al., 1964a, b; Tundisi and Krishnaswamy, 1967; Omori, 1969a; Tracy and Vallentyne, 1969; Vinogradov et al., 1970; Clutter and Theilaoker, 1971; Childress and Nygaard, 1974; Salonen et al., 1976.

ranges. Bordovskiy et al. (1976) estimated 65.2% of the dry weight of Gnathophausia gigas as carbon, a value somewhat higher than those given in Table XII. Jawed (1969) found 8-8-11.4% dry weight of Neomysis rayii as nitrogen. Some 1 6 2 3 % of the total nitrogen in Neomysis integer was found to be non-protein nitrogen (NPN), an increasing concentration within this range being associated with increasing environmental salinity ; this NPN is predominantly present &S free amino acids (Raymont et al., 1968). A lower range of NPN,

102

THE BIOLOGY OF MYSIDS

7-10% of the total nitrogen, was recorded in the warm-water species Taphromysis bowmani by Johnson and Hopkins (1978). Mysids, unlike euphausiida, have seldom been used in studies of uptake, retention and dispersion of radioactive isotopes. Fukai et al. (1962) recorded 0.8 p Ci90 Sr/kg wet weight in Neomysis japonica in Tokyo Bay in June, 1959. Marshall et al. (1975)studied the partitioning of plutonium-239, originating from radioactive fallout from weapons’ testing in the Great Lakes, and found that Mysis relicta and the amphipod Pontoporeia afinis Lindstrom had higher levels present in their bodies than other crustaceans. The concentration factors for Plutonium in these two species are 760 and 1600 respectively (Yaguchi et al., 1974). The higher concentrations of plutonium in these animals were assumed to have arisen from their feeding close to the surface of the sediment where the highest environmental accumulations of the isotope occurred. Marshall et al. (1975) also examined the ratios of fallout plutonium-239 to caesium-137 in the animals and environment during this period, 1972-1973. The calorific value of Mysis stenolepis has been estimated and found to range from 4563 to 4824, mean 4714, gcal/g dry weight representing a mean value of 99Ogcal/g wet weight (Tyler, 1973). The calorific values of temperate and high latitude species are likely to show seasonal variation as Tyler has demonstrated in several species of organisms, and Brattelid and Matthews (1978) showed in Boreomysis arctica. The calorific value of some deep-sea species has been calculated by Childress and Nygaard (1974) who found a range of 400-1500 g cal/g wet weight in four species of Gnathophausia and in Boreomysis californica. Johnson and Hopkins (1978) found that female Taphromysis bowmani have calorific contents consistently higher than males; the ranges were 4428-4712 g cal/g dry weight in females and 4178-4227 g cal/g dry weight in males. A similar range of values was found in Neomysis mirabilis by Shushkina et al. (1971). The calorific content of male Mysis relicta showed a loss of about 8% over the period of the die1 vertical migration (Teraguchi et al., 1972), the mean content being about 6200 g cal/g dry weight; such a loss was not detected by Barker (1973) and Foulds and Roff (1976), the former recording a range of 5500-7000gcal/g dry weight in this species. Wissing et al. (1973) measured a calorific value of 7533 g cal/g dry weight in unidentified mysids from the Gulf of Mexico. The calorific content of Boreomysis arctica increases slightly with increasing body weight in maturing and spent females but decreases slightly with increasing body dry weight in males. Males have a lower calorific content than maturing females but small spent females have the least

6.

103

CHEMICAL COMPOSITION

calorific content of all (Brattelid and Matthews, 1978). Maturing females, spent females and males of 15 mg body dry weight have calorific contents of 6500, 5975 and 5830 g cal/g dry weight respectively. Carbohydrates occur a t low concentrations in mysids (Table X, p. 99. Raymont et al. (1968)) in preliminary experiments, concluded that 4&50% of the carbohydrate in N e o m y s i s integer is present as insoluble material, probably as ribose (as nucleic acids). Some soluble pentose, about 40% of the total carbohydrate, is also present. Only traces of glucose and galactose have been detected. The glycogen stores appear to be utilized rapidly under conditions of unsatisfactory feeding, for example when the animals are caught in the wild and transferred to laboratory conditions ; the largest concentrations of glycogen were recorded from freshly caught individuals. The amount of information available about the lipids of mysids is less than that for the lipids of euphausiids. L. R. Fisher determined the gross lipid content of a number of species and his unpublished data are presented in Tsbles XI11 and XIV. Further analyses from TABLEXIII. LIPIDS(yoWET WEIQHT)IN SPECIESOF LOPHOGASTRIDA (L. R. Fisher, unpublished) Average body weight Species Eucopia grimaldii E . sculpticauda

Gnathophawria gigas G . gracilis

G . iagens

G. zoea

Lophogaster spinoswr L. typicwr

Sea area

Date

N. Atlantic N. Pacific N. Atlantic N. Atlantic N. Atlantic N. Pacific N. Pacific N. Atlantic Madeira N. Atlantic Madeira N. Pacific Madeira N. Atlantic Madeira Madeira N. Atlantic N. Atlantic N. Atlantic N. Atlantic

November 1955 April 1956 November 1955 November 1957 April 1958 April 1956 April 1956 November 1955 July 1954 November 1954 May 1955 April 1956 June 1956 October 1959 May 1955 June 1956 April 1958 October 1959 November 1955 September 1955

yo

(ms)

lipid

60 171 146 295 258 786 924 1964 11152 446 11536 3812 6167 3703 1108 733 571 374 70 2.4

8.1 24.0 12.0 12.0 15.0 3.3 12.0 26.0 6.4 2.5 5.2 9.4 2.1

8-9 14-0 1.9 15.0 7.4 9.7 8.3

104

THE BIOLOGY OF MYSIDS

TABLEXIV. LIPIDS(% WET WEIUHT) IN SPECIES OF MYSIDA (L: R. Fisher, unpublished)

Species Acanthomysis longicornis Amblyops kenvpi Boreomysis arctica B. californica Erythrops erythrophthalma Ewhaetomera tenuie Gaatrosaccwr spinifer Hemimyeis lamornae Leptomysis gracilis

L. mediterranea Mysidopsis didelphys

Neomysis americana Paramyeis arenosa Schistomysis ornata Siriella armata s. clawrii S. norvegica

Date

Average body weight fmg)

N. Atlantic

November 1955

5.3

1.9

N. Atlantic N. Atlantic N. Pacific Barents Sea

July 1955 September 1955 April 1956 January 1955

21.0 8.3 35.0 3.8

6.1 4.8 4.8 4.3

N. Atlantic N. Atlantic Plymouth N. Atlantic Scotland Monaco Scotland Scotland Scotland Scotland Scotland Scotland Scotland Scotland Virginia, USA Monaco Scotland Scotland Scotland Monaco N. Atlantic Scotland Scotland

September 1955 September 1955 June 1952 September 1955 November 1955 February 1952 June 1954 September 1954 September 1954 November 1954 December 1954 March 1955 November 1955 November 1959 February 1954 February 1952 March 1955 November 1955 April 1957 February 1952 November 1955 December 1955 March 1955

4.8 1.1 5.0 5.5 4.6 4.9 28.0 20.0 19.0 19.0 18.0 20.0 18.0 21-0 27.0 2.2 9.0 10.0 12.0 22.0 1-1 93.0 7.1

Sea area

yo lipid

3.3 20-0 3.4 0.8 13.0 1.5 6.2 4.1 3.7 2.6 2.2 7.1 1.3 7-8 0.5 8.6 1.8 0.3 0.6 1.9 6.0 4.0 5.6

other sources are given in Table X and XV. The highest values for lipids, with a modal range of 5-12% of body wet weight, occur in species of the genera Gnathophusia, Lophogaster and Eucopia .of the Sub-order Lophogastrida. Lower values, with a modal range of 1-6% of body weight, occur in species belonging to the genera of the SubOrder Mysida. Species of the Lophogastrida are predominantly mesoand bathypelagic, but some of the species of Mysida enalysed are also deep living, for example Boreomysis arctica, B. californica and

6. CHEMICAL COMPOSITION

105

TABLEXV. LIPIDS(yoWET WEIGHT)IN MYSIDS Species Qnathophawia gigas ffnathophawiasp.

Lophogaster sp. Eucopia sculpticauda males females Neomy& integer gravid females females males juveniles N . awatschensis N . intermedia N . nakazawai

N . spinosa

Sea area

Date

yo lipid

Kuril-Kamchatka 31"N 119OW 29"N 23"W 30"N 23OW 37"N 14OW 37"N 14"W 30"N 23"W 25"N 19"W

Summer April March April April March November

63= 42a 1.0 2.3 4.7 1.2 0.7 1.7

37"N 25"W 37"N 25"W

October October

9.7 11.5

England England England England Japan Japan Japan

October October October October -

Japan

-

-

1.7 1.6 0.9 1.1 1.3 3.9" 6*5a 1.34 (10.3a) 2.0

Data from: Yamada, 1961, 1964; Lee et al., 1971; Morris, 1972, 1973; Morris and Sargent, 1973; Bordovskiy et al., 1976. a On dry weight basis.

Amblyops kempi. There would, therefore, appear to be a relationship between increasing depth of occurrence and increasing lipid storage but further data are required to confirm this. Childress and Nygeard (1974), in a general study of the chemical composition and buoyancy of midwater organisms, found that lipid contenb increased with increasing depth but decreased again at the greatest depths. Female Mesopodopsis slabberi and Neomysis integer store significantly greater quantities of lipids than males. Consequently, the sex ratio of mysids in samples will affect the estimations of lipid content, not only in these two species but probably also in most other species. Seasonal variation in the lipid content of euphausiids has been demonstrated in temperate and high latitude species. Moore (1976), however, found relatively constant levels of lipids present in the mysids Neomysis integer and Praunus JEexuosus, although he did not analyse samples in the autumn and early winter. Linford (1965), although finding fluctuating levels of lipids, did not demonstrate a

106

THE BIOLOQY OF MYSIDS

significant seasonal variation in their concentrations in Mesopodopsis slabberi. Such fluctuations are present in Fisher’s data on Mysidopsis didelphys (Table XIV) ; a seasonal trend could be inferred were it not for the high value of 7.8% in November, 1959. A continuing decrease in the lipid content has been shown to occur throughout the summer in Neomysis integer by Morris (1971). Animals that he kept a t different temperatures in the laboratory showed a correlation between higher lipid concentrations and lower environmental temperatures. Bamstedt ( 1 978) examined the seasonal fluctuations in protein, carbohydrate, lipid, chitin and ash in Boreomysis arctica but expressed all his data as percentage dry body weight. He determined the seasonal changes in the dry weight of two standard mysids, one of 19 mm and the other of 13 mm total body length, over a period of 13 months. The dry weight content was highest in the autumn and early winter when the animals were preparing to reproduce. These changes in the dry weight undoubtedly reflect changes in the lipid content. Seasonal fluctuations in the quantities of lipids stored are expected to occur in many species but such fluctuations have not yet been clearly demonstrated. Some general characteristics of the lipids are given in Tables XVI and XVII. Yamada (1964) examined some properties of the lipids of Neomysis mirabilis, N . intermedia, N . spinosa and N . awatschensis. Their density ranged from 0.9407 to 0.9561 and their refractive index from 1.4639 to 1.4866. The lipids can be divided into various fractions such as monoglycerides, diglycerides, triacylglycerols, phospholipids, free fatty acids, wax esters, sterol esters and sterols (Sargent, 1976). The dominant fractions within the lipids are the phospholipids, triacylglycerols, free fatty acids and, in some species, wax esters. The proportions of some of these fractions in the lipids of several species are shown in Table XVII. These data demonstrate the virtual absence of wax esters in shallow living species, represented in the genera Neomysis and Praunus, and their dominance in the lipids of the offshore, deep living lophogastrid species in the genera Gnathophausia, Lophogaster and Eucopia. Sargent et al. (1978), studying several species of mesopelagic decapods and Eucopia sculpticauda, concluded that the wax esters are largely produced by the biosynthetic activities of the animal’s metabolism rather than supplied to it in its diet. The fatty acid composition of the total lipids of several species are shown in Table XVIII. Lewis (1967) examined the lipids of Gnathophuusia sp. and found a high proportion of C-18 acids. The major fatty acid and fatty alcohol constituents of the wax esters are C-18: 1 acid and C-16: 0 alcohol (Table XIX). Morris (1972) found 51-58% of the wax esters to be oleic acid (C-18: 1) and 67-73% of the wax

TABLEXVI. CHARACTERISTICS OF LIPIDSOF Neomysis SPECIES (Yamade,1964; Raymont et al., Acetone soluble

Nonsapon.

(%I

di6

ng

sv

IV

AV

(%I

N . awatachensis N . integer N. intediu N . mirabilis

95.8

0.9407

1.4838 1.4821 1.4639

195.7 137.0 109-8 181.3 200.4 202.6

116.8

0.9561

175.2 143.0 150.0 163.6 186.9 179.5

11.3 14.0 17.89 10.62 11.12 10.10

98.5 96.9

acid

Bromides insol. in ether

IV

(%I

Faty

Species

N . spinosa

1968)

0.9468 1.4866

76.4 133.2 90.4 76.0

NV

69.55 197.6 190.0 193.2 195-2

121.5 207-9 192.0 208.1

dC,density; n$, refractive index; SV, saponification value; IV,iodine value; AV, acid value; N V , neutralization value.

26.75 73.7 79-92 74.95

108

THE BIOLOOY OF MYSIDS

TABLEXVII. PERCENTAGE COMPOSITION OF TOTAL LIPIDS Species Gnathophazcsia sp.

Lophogaster sp. Eucopia sculpticauda males females Praunw Jexuoaua N e m y s i s integer

Phospholipida

Triacylglycerole

11 11 6 18 18

26 ( + F F A " ) 35 11 44 37 (FFAO) 12

Wax esters 60 52 77 32 40 69 5 67 77

24-27.5 50 2 3-2 7.5

34 6gb

0.90

Date from: Raymont el al., 1968; Lee et al., 1971; Morris, 1972; Lee and Hirote, 1973; Morris and Sargent, 1973; Morris et al., 1973; Moore, 19.76. Plus free fatty acids. Plus diglycerides. Plus sterol esters.

alcohols to be (3-16: 0 alcohol in male and female Eucopia sculpticauda. Species of Gnathphawia have been shown by Morris and Sargent (1973) to be able to synthesize the C-18: 1 acids and C-16: 0 alcohols in the wax esters, as well as the higher polyunsaturated fatty acids, from palmitic acid. Similar biosynthesis of these long-chain fatty acids for incorporation in the phospholipid and triacylglycerol fractions of the lipids has been demonstrated in Neomysis integer by Morris et al. (1973). Further, N . integer was able to synthesize a significant portion of its required lipid directly from starch but after a week on a starch diet the levels of long-chain polyunsaturated fatty acids declined (Morris et al., 1977). The proportions of the constituent fatty acids of the lipids fluctuate. Morris (1971) showed, in N . integer, that the polyunsaturated acids increased during the summer and autumn while the short-chain low unsaturated acids decreased. Similarly, he found that an increase in the long-chain polyunsaturated C-22: 6 acids took place at the expense of the short-chain low unsaturated acids when environmental temperatures decreased. In addition to such variations, Morris (1973) demonstrated sexual differences in the fatty acid compositions of male and female N . integer, with juveniles somewhat intermediate in composition. The females had a higher total lipid content associated with a

TABLEXVIII. FAW ACIDCOMPOSITION OF Lmms OF M Y S ~ (%) S

Patty acid 10 : 0 12 : 0 13 : 0 14 : 0 15 : 0 16 : 0 17 : 0 18 : 0 19 : 0 21 : 0

Total

Neomysis integer

Neomysis awatschensis

Neomysis mirabilis

Anchialina agilis

Lophogaster

Tr

Tr

0.4

0.2

0.3

0.5

0.3

0.2

Tr

Tr

Tr

Tr

3.8 0.8 21.3 1.9 5-5 0-7 0.4 34-6

4.6 18.2 21.2 1.6 3.0

6.9 0.8 20.1 0.9 3.3 0.4 0.5 33.3

4.7 1.2 20.9 1.4 4.4 0.5

5.3 1.6 24.5 1.1 3.8 0.4

33-4

36-9

7.4

0.5

0.8 0.4 10.5 1.5 15.7

5-1 0.3 13.2

7.0 0.9 13.4

11.5 1.3 50-6

0.4

0.7

7.5

19.0

22-0

70.9

Tr 3.9 0.7 25.2

0.8 4.0 0.5 0.5 36.0

Tr 31.9

Neomysis spinosa

8P.

Gnathophausia SP.

0.8 0.3 6.3

Tr

Mono-unsaturated 14: 15: 16: 17: 18: 19: 20: 22: 24:

1 1 1 1 1 1 1 1 1

Tote1

10.3 0.6 12.4

10-9

0.6

1.3 1.7

Tr

Tr

23.9

27.7

31.1

13.3

2.2

0.6 10.4 11.6 1.6 1-6

Tr 25.8

cod.

TABLEXVIII-contd.

Polyunsaturated 16 : 2 16 : 3 18 : 2 18 : 3 18 : 4 20 : 2 20 : 3 20 : 4 20 : 5 22 : 4 22 : 5 22 : 6 Total

Tr 0-2 0.7 0.4 0.3 1.7 21.5

Tr

3.0 3.4 1.5 2.3 0.9 Tr 17.5

1.8 1.0 2.8

1.5 0.6 2.6 Tr 0.7 Tr 21.1

Tr 24.2

Tr 11.4 37.0

Tr 10.5 40.0

9.1 39.9

Data from: Yemade, 1964; Lewis, 1967; Morris, 1971.

Tr

0.8 0.7 0.9 Tr

3.7 1.6

0.5

Tr

1.0

0.5

Tr 14.7 40.0

1.0 0.8 0.8 0.2 2.6 16.2 Tr 2.1 22.9 46.6

2.0 12.1 1.0 0.2 23.0 40.7

4.8

5-9

6.3 22.8

6.

111

CHEMICAL COMPOSITION

TABLEXIX. FATTY ACID COMPOSITION OF LIPIDFRACTIONS

Fatty acid

Neomysis Praunus integer flexuosus (yomolar concentration)

Gnathophausia 8p. (fatty acid and fatty alcohol) (yocomposition) ~~

PL 14 : 0 15 : 0 16 : 0 16: 1 16 : 2 17 : 0 l? :1 18 : 0 18: 1 18 : 2 18 : 3 18 : 4 20: 1 20 : 4 20 : 5 22 : 1 22 : 6 22 : 6 Unidentified

4.3 0.6 26.1 9.2 0.1 2.3 20.6 1.1 0.8

TG 6.9 0.5 21-3 4.5 4.3 4.2 14.7 1.5 0.5

PL 3.9 25.1 6.9 0.4

3.6 23.4 0.6 1.7

TG 5.3 0.2 23.2 5.1 2.3

4.9 20.7 2.0 1.1

21.3

25.3

19.3

22.1

13-1 9.5

9.8 6.5

14.2 0.9

11.4 1.7

PL 1-7 0.8 15.9 3.9 0.7 3.1 1.1 17.3 1.4 1.2 1.0 Tr 5.1 14.3 Tr Tr 32-5

TG WE" WEb 2.3 0.4 4-6 0.9 16.5 2.0 82.6 6.6 10.7

Tr 4.0

2.3 31.7 1.3 0.6 2.0 2.0 9.9 2.3 Tr 17.5

73.7

10.2 2,5

8.6 0.6 3.7 0.3

PL, phospholipids: TG, triacylglycerols; WE, wax esters. Data from: Morris and Stlrgent, 1973; Moore, 1970. acid. alcohol.

larger triacylglycerol fraction. The larger triacylglycerol fraction resulted in increased levels of mono-unsaturated acids in the total lipids of the females which were further found to have reduced levels of C-22 : 6 polyunsaturated acids, when compared with the compositions of males and juveniles. Sterols have noti been studied to any degree in mysids. Morris (1973) found 4.8% of the lipids of N . integer as sterols. Cholesterol and provitamin D were identified in the sterols extracted from a mixed sample of Mysis microphthalma and Paramysis loxolepis by Vinogradova and Kandiuk (1967). Teshtimrt and Kanazawa (1971) found that the concentration of sterols in unnamed mysids was 0.21y0 of the body wet weight; this sterol mixture consisted of 3% as 22-dehydrocholesterol, 78% as cholesterol, 8% as brassicaterol, 10% as 24-methylenecholesterol, and a trace of B-sitosterol.

112

THE BIOLOGY O F MYSIDS

The amino acid composition of the protein hydrolysates of Neomysis integer and Mesopodopsis stubberi has been investigated and the predominant acids, in descending order of importance, found to be glutamic acid, aspartic acid, lysine and arginine (Table XX). There is a close similarity between the amino acid composition of these species and that of the freshwater Mysis relicta from Cayuga Lake, New York (Table XX). The most important free amino acids are glycine, taurine, and arginine (Table XXI). The general pattern of amino acids present in the two species are closely similar. Raymont et al. (1968, 1973) and Armitage et at. (1977, 1978) discuss the factors producing variation in the concentrations of individual acids. They also examined transamination between 17 amino acids and the keto acids, a-keto-glutarate, pyruvate, and glyoxylate in Neomysis integer. TABLEXX. AMINOACID COMPOSITIONOF PROTEIN HYDROLYSATES OF MYSIDS (yoOF TOTAL) Neomysis integer

0uig erow Amino acids

Cysteic acid Taurine Aspartic acid Threonine Serine Glutamic acid Proline GI ycine Alanine Valine Cystine Methionine Isoleucine Leucine Tyrosine Phenylalanine Lysine Histidine Tryptophan Arginine Ornithine Total (mg/g dry weight) yo dry weight

females

11.2 4.1 3-8 17.2 3.0 3.6 5.2 5,2 1.1 2.3 5.3 7.7 4.0 5.1 10.2 3.5 7.3 0.2 464.9 46.5

Less mature

Mesopodopsis slabberi

10.6 3.9 3.5 19.1 2.4 4.1 5.4 5.4 0.7 1.7 5.3 8.4 3.9 5.0 10.2 2.4

-

7.8 0.2 510.1 51.0

Data from: Raymont et al., 1973);Tracy and Vallentyne, 1969.

11.0 4.4 4.6 14.8 3-2 4.0 5.8 5.6 1.4 2.4 5.3 7.8

4.3 4.6 10.4 3. ” 6.7 0.2 548.4 54.8

Mysis relicta

11.7 5.4 6.1 14.7 3.8 7.4 8.0

5.5 0.8 1.7 4-9 9.1 2.1 3.9 7.5 1.3

-

5.7

6, CHEMICAL COMPOSITION

113

TABLE XXI. FREEAMINO ACIDSOF MYSIDS (yoOF TOTAL). ONLY EIGHTOF THE MAJORACIDSARE LISTEDIN THREESETSOF DATA Neomysis integer Anzino acids

Cysteic acid Taurine Aspartic acid Threonine Serine Glutamic acid Proline Glycine Alanine Valine Cystine Methionine Isoleucine Leucine Tyrosine Phenylalanine Lysine Histidine Tryptophan Arginine Ornithine Total (mg/g dry weight) yo dry weight

Neomysis integer 0-8 23.1 1.9 1.3 1.4 5.5 4.2 30.9 3.8 1.0 0.6 0.2 0.6 0.7

Owigerow females

Lem mature

Mesopodopsis slabberi

26.4 3.1

26.6 2.9

14.7 4.7

6-0 5.9 27.4 4.8

5.6 2.4 29.6 5-2

22.5 2.9

1.3

2.2

9.0

17.6

12.0

25-5

7.9

-

0.6 0.4 1.5 1.5 0.7 19.1 0.1 95.7 9.6

~~

Data from: Srinivasagm et al., 1971; Raymont et at., 1973.

The a-keto-glutarate transaminase reaction was strong but a weak glyoxylate transamination was also present suggesting some direct production pathways for the free amino acid glycine. Vitamin B,, was found in Neomysis intermedia at concentrations similar to those in euphausiids and markedly higher than in the other crustaceans examined (Hirano et al., 1964b). The occurrence of vitamin C was examined in mysids by Santa and Bacescu (1942). They found the following quantities, in rng/100 g wet weight, near Roscoff, France : Leptomysis lingvura, males and females, 0.14; Mesopodopsis slabberi, males and females, 0.10; Neomysis integer, males and non-ovigerous females, 0.06-0.07, ovigerous females, 0.09 ; Paramysis arenosa, juveniles 0.30, males, 0.21 and L,

114

THE BIOLOGY OF MYSIDS

ovigerous females, 0.13 ; Praunus Jlexuosus, males, 0.1 1, females, 0.13 ; Schistomysis parkeri, ovigerous females, 0.20; S. spiritus, males and females, 0.15. Vitamin A occurs in some species of mysids and not in others. L. R. Fisher (see Mauchline and Fisher, 1969) studied its occurrence in euphausiids where it is predominantly located in the eyes. He published the results of analyses of a few species of mysids (Fisher et al., 1953; Fisher and Kon, 1959); other unpublished data are included in Table XXII. No vitamin A was found by him in the following species : Amblyops kempi, Anchialina agilis, Boreomysis arctica, B. microps, Erythrops erythrophthalma, E. serrata, Euchaetomera tenuis, Eucopia grimaldii, Gastrosaccus spinifer, Hemimysis lamornae, Leptomysis mediterranea, Lophogaster spinosus, L. typicus, Meterythrops picta, Paramysis arenosa, Xiriella armata, S. clausii, S. jaltensis and S . norvegica. Vitamin A was not detected in one sample of Gnathophausia zoea, as shown in Table XXII. Fisher concluded from his studies on Crustacea that pelagic species were richer in vitamin A than neritic or benthic species; there ia no clear evidence of this in mysids. His data show that more vitamin A was present in the eyes than in the bodies of only four species of mysids-Gnathophausia zoea, Eucopia sculpticauda, E . unguicubta and Mysidopsis didelphys (Table XXII). The concentration of vitamin A shows seasonal changes in euphausiids and this may also be true in some species of mysidsj the data for Mysidopsis didelphys (Table XXII) suggest the occurrence of high concentrations in the winter to spring period. Carotenoids were first identified in mysids by Lonnberg (1931a, b, 1934) who examined Praunus Jlexuosus. Fisher estimated the concentrations of astaxanthin in a few species (Table XXII); in addition to these, he found that Leptomysis mediterranea had 39 pg, Paramysis arenosa 37 pg and Xiriella armata 16 pg astaxanthinlg wet weight. These concentrations are within the ranges occurring in euphausiids (Mauchline and Fisher, 1969), except for the t w o very high values found in Erythrops erythrophthulma which were sampled in the Barents Sea, and Mysidopsis didelphys sampled in March in Loch Fyne, Scotland. Seasonal fluctuations in concentrations are suggested by the data on M . didelphys (Table XXII). The pigments of mysids have received little attention. Fisher (unpublished) examined the eye pigments of Eucopia sculpticauda and found evidence suggesting that the cone pigment is melanin and that the inner pigment surrounding the rhabdoms is also melanin. Elofsson and Hallberg (1973), however, state that the black pigment in the eyes and integument of Neomysis integer is an ommochrome.

6. TABLEXXII. VITAMIN A

Species

115

CHEMICAL COMPOSITION AND

ASTAXANTHIN (L. R. Fisher, unpublished)

Vitamin A (pg/g wet wt)

Gnathophausia gigas G. gracilis (I.ingens

G. zoea Eucopia sculpticauda E. unguiculata Boreomysis calif ornica Boreomysis sp. Erythrops erythrophthalma Katerythrops sp. Leptomysis gracilis Mysidopsis didelphys (March) (June) (September) (September) (November) (December) Schistomyeis ornata

% in eyes

3.1 1.7 0.9 1.7 0.6 2.0 0.2 0.9 1.4 0 1.3 1-7 0.2 0.3 3.0 0.9 0 0.9 0.6 0.7

0 19 30 47 0 4.4 13 100

6.5 3.2 0.8 0.8 0.6 0.8 0.3

100 100 50

Astaxanthin ( w / qwet wt)

57 60 76 87

Habit" Bathypelagic 1000-2000 m 350-400 m

400-2000 m

100

850-2600 m

54

700-2500 m

10

Bathypelagio 125

40-500 m 5-500 m

100 100 0

112 29 63 64 42 31

60-300 m

50-150 m

0.5

Neomysis americana Acanthomysis longicornis

0.7 1.6

0-240 m

< 100 m

Mauchline end Murano (1977).

The chromatophores of N . integer and Praunus JEexuosus contain many microtubules and several chromatophores of different colours unite to form chromatosomes. Some contain granules characteristic of red carotenoid pigments. Three white pteridines, dominated by isoxanthopterin, probably constitute the reflecting pigments of the eyes. The chemical nature of the yellow pigment that is also present in mysids remains unknown. The haemolymph of Gnathophausia ingens has an absorption spectrum that suggests the presence of a haemocyanin (Freel, 1978).

116

THE BIOLOGY OF MYSIDS

Succinic dehydrogenase activity was found to be inversely proportional to body weight in Neomysis integer by Raymont et al. (1967). Temperature accelerated the activity and there was evidence of seasonal variation. Adenine nucleotides have been examined in Boreomysis arctica and Hemimysis abyssicola by Skjoldal and Bamstedt (1977). A seasonal fluctuation in adenosine triphosphate (ATP) concentration was observed, a maximal concentration of 6-7 pg ATP/mg dry body weight being present in April-May in the Norwegian populations. Comparable concentrations of adenosine diphosphate (ADP) and adenosine monophosphate (AMP) were 3-4 and 1-3 pg/mg dry body weight respectively. Homarine, a betaine that is supposed to have an osmoregulatory function, was identified in Neomysis intermedia by Hirano (1975). It is N-methyl picolinic acid betaine and occurred at a concentration of 0.58 mg/g wet body weight, a concentration equivalent to about one fifth of that present in the euphausiid, Euphuusia pacijica.

CHAPTER I

INTERNAL ANATOMY The internal anatomy of a few species of mysids, such as Praunus jlexuosus and Gnathophausia species, has been examined in some detail. The structure of the male and female reproductive system has already been described, while discussing the marsupium, and the histology and form of the alimentary tract is described in the section on feeding. Information on other aspects of the internal anatomy is reviewed here. Delage (1883) describes the circulatory system of “Mysis” and his diagrams suggest that the species was Praunus Jlexuosus; it is certainly a species of the tribe Mysini. Siewing (1956) describes the system of Eucopia sculpticauda and E . unguiculata. Mayrat (1956b) examines the blood supply to the head while Alexandrowicz (1955) studies the nervous innervation of the heart of P . jlexuosus. More recently, Belman and Childress (1976) have examined the circulatory system of the lophogastrid mysid, Gnathophausia ingens. The heart is a single chamber, tube-like in form and situated in the posterior dorsal region of the thorax. It is suspended within the pericardium (Fig. 30). The heart is relatively large, Belman and Childress (1976) finding that its wet weight represents 0.20-0.32% of the wet body weight of G. ingens. Valves are present at each end and mark the extent of the heart because the “tube” continues anteriorly as the aorta cephalica and posteriorly as the arteria abdominalis (Fig. 30). Paired ostia, a left and a right, are located dorso-laterally in the median region of the heart; blood enters the heart through them from the pericardium. There are three systems of nerve elements in the heart of Praunusflexuosus (Alexandrowicz, 1955). The first is a local system situated on the outside of the dorsal wall of the heart, the second consists of the nerves connecting this local system to the central nervous system, and the third is the nerves of the arterial valves. The heart first, starts to beat in the Stage I1 larva within the marsupium but beating is discontinuous, occurring in rhythmic bouts lasting a few seconds. Berrill (1969) found that the bouts would occur every 3-4 min a t 4°C and every 1-2 min at 15OC in larvae of Mysis relicta.

118

THE BIOLOGY OF MYSIDS

f

C FIG.30. The circulatory system of mysids. A and B, the system in a species of the tribe Mysini. C, the system of Gnathophausia ingens. The arteries are shown in black; the pericardium is stippled in B and C and the afferent and efferent vessels and gills are indicated in C by dotted vessels. A and B are constnicted from figures of Delage (1883) and Mayrat (1956b); C is after Belman and Childress (1976). aa, arteria abdominalis; ab, arteries to the cerebral ganglia; ac, aorta cephalica; ae, afferent and efferent blood vessels of gill; al, left arterk lateralis; an, antenndar and antenna1 arteries; e, efferent branchial channel; fh, frontal heart; g, gill; h, heart; ha, hepatic arteries; hal, left hepatic artery; har, right hepatic artery; oa, optic artery; p, pericardium; a,sternal artery; sal, left sternal artery; sar, right sternal artery; ua, arteries of unknown destination; va, ventral abdominal artery.

The heart commences beating continuously in the late Stage I11 larva, immediately before it emerges from the marsupium. Henderson (1927), working with what he calls Michtheimysis (probably Mysis relicts), found the heart of the adults beating a t a rate of 120-380 beatslmin, over a temperature range of - 2-18°C; there was a linear relationship

7. INTERNAL ANATOMY

119

between increasing rate of heart beat and increasing temperature. Belman and Childress (1976) found that larger Gnathophuusia ingens have a heart rate varying from 128 to 178 (mean 110) beatslmin a t 4OC and a t 1 atmosphere hydrostatic pressure. The heart rate of G . zoea was approximately 60 beatslmin a t 5°C and 200 atmospheres pressiire during compression experiments made by Macdonald and Gilchrist (1978). The arterial system of mysids is shown in Fig. 30 but some of the descriptions appear to be contradictory and further investigation is required. The aorta cephalica travels anteriorly round the dorsal side of the stomach t o the head supplying small branches to the intestine, cardiac stomach and various muscles (Belman and Childress, 1976). It then divides, a branch descending ventrally to the cerebral ganglia. A frontal heart is located antero-dorsally to this branch (Sewing, 1966; Mayrat, 1956b; Belman and Childress, 1976). This is a swelling of the aorta cephalica associated in Praunus jexmsus with the division of the artery into the branch to the cerebral ganglia and those to the eyes. The optic arteries, however, do not arise from the aorta cephalica in Gnathphausia ingens (Belman and Childress, 1976) and this is surprising because the eyes are supplied by this artery in other mysids, and in euphausiids (Mauchline and Fisher, 1969) and in some decapods (R. S. Pillai, 1965). The frontal heart contracts rhythmically, according to Belman and Childress, more or less in phase with the heart. Ventral to the optic arteries, the aorta cephulica supplies the antennules and antennae in Praunus jexuosus (Delage, 1883) but not in Gnathophuusia ingens (Belman and Childress, 1976). Delage (1883) figures paired hepatic arteries a t the site of the anterior end of the heart where Belrnan and Childress figure paired arteriae laterales. The arteriae laterales in G . ingens supply small branches to the digestive glands, anterior regions of the stomach and various muscles and then supply the eyes, antennules and antennae. These arteries supply the antennae, but not the eyes, in euphausiids and other decapods. In the case of Praunusjexuosus (Fig. 30) it is possible that Delage has wrongly named the arteriae laterales as hepatic arteries. The hepatic arteries originate in the region between the arteriae laterales and the anterior end of the heart in Gnathphuusia ingens (Fig. 30), in euphausiids (Mauchline and Fisher, 1969) and in decapods (R. S. Pillai, 1965). Delage (1883) figures two arteries in this region (Fig. 30) going to unknown destinations; Alexandrowicz (1955) mentions these arteries of Delage, but provides no further information. These arteries of Delage could be the true hepatic arteries of Praunus jlexuosus.

120

THE BIOLOGY OF MYSIDS

The aorta descendens or sternal artery originates at the posterior end of the heart. It is single in PraunusJlexuosus (Fig. 30),in Eucopia species (Siewing, 1956), in euphausiids (Mauchline and Fisher, 1969) and in decapods (R. S. Pillai, 1965) but paired in Gnathophausia ingens (Fig. 30). An anterior ventral branch supplies the ventral region of the thorax, thoracic legs and maxillae. The more anterior mouthparts are probably supplied by the arteriae laterales. A posterior branch of the aorta descendens (sternal artery) is shown by Delage as not extending into the abdomen in Praunus Jlexuosus and similarly by Siewing in Eucopia species, but described by Belman and Childress as extending posteriorly as the ventral abdominal artery in Gnathophausia ingens (Fig. 30). Dorsal and lateral regions of the abdomen are supplied by the arteria abdominalis which terminates in branches to the telson and uropods. The lateral branches of these arteries also supply the pleopods. The various arteries sub-divide into arterioles within the tissues that they supply and the blood is voided from their ends into a complex system of sinuses. Belman and Childress briefly describe the form of the sinuses in G. ingens. Blood from the anterior thoracic sinus and from the abdominal sinuses travels to the infrabranchial sinus. It then enters the afferent branchial channels and so into the afferent branchial veins within the gills. There it circulates, returning through the efferent branchial veins out of the gills into the efferent branchial channels which lead into the pericardium (Fig. 30). Once there, it enters the heart through the paired ostia which are closed by valves during the systolic contraction, Praunus Jlexuosus, like all non-lophogastrid mysids, has no gills. Delage concludes that oxygenation of the blood takes place in the sinuses under the carapace. There are various marked differences between the circulatory systems described in Praunus jlexuosus and Gnathophausia ingens noted in the above account. These may be real differences and R. S. Pillai (1965) found such differences existing between descriptions of the circulatory systems of several decapod crustaceans. Elucidation of the circulatory system is difficult, especially the tracing of the destinations .of the smaller arteries. There does appear, however, to be a general pattern of blood vessels common to mysids and decapods. This basic pattern is also evident in the arterial system of the gammarid, Gammarus pulex (Cussans, 1904) and the isopod, Ligia oceanica (Hewitt, 1907). Consequently, further investigation of the circulatory system of mysids may confirm that there are errors in the present descriptions. The blood pressures in various regions of the circulatory system of Cnathophausia ingens have been measured by Belman and Childress

7.

INTERN&

ANATOMY

121

(1976). The maximum intracardiac systolic pressure was 27.0-20.5 cm H,O with a mean vhlue of 24cm H,O. The range of diastolic pressures was 21.0 to 16.0 cm H,O with a mean of 19 cm H,O. High arterial pressures, almost equal to that of the heart at systole, are maintained in the aorta cephalica and arteria abdominalis. The blood velocities had a mean systolic value of 2-71 cmjs. Osmotic regulation of the blood concentration takes place in brackish water species of mysids. Hyperosmoticity has been demonstrated in low environmental salinities and hypo-osmoticity in 'high environmental salinities (Khlebovich et al., 1970; McLusky and Heard, 1971; Dormaar and Corey, 1973). The iso-osmotic point for Praunus Jlexuosus was at 25%,, while that of the more brackish living species Neomysis integer was at 19X0 (McLusky and Heard, 1971). Another coastal species, Mysis stenolepis, had an iso-osmotic point a t 2 l%,,(Dormaar and Corey, 1973). Mysis relicta, collected from Canadian freshwater lakes, hy-perosmoregulates a t salinities below about 25%, and is isoosmotic to its environment at salinities between 25 and 30X0 (Dormaar and Corey, 1978). G. A. Vinogradov (1976) compares the ability of M . relicta and M . oculata to osmoregulate. The concentrations of sodium, potassium and chloride ions in the haemolymph are isoionic to dilute sea water but hypo-ionic to full strength sea water. The abdominal muscular systems of Gnathphausia zoea, Lophogaster typicus and Praunus flexzlosua are described by Daniel (1928, 1931, 1932, 1933) and that of Spelaeomysis longipes by Nath (1974). Manton (1928a) compares the musculature of the fifth and sixth abdominal segments of Gnathophausia zoea, Lophogaster typicus, L. interrnedius and Hemimysis lamornae. The general arrangements of the muscles are shown in Figs 31-33. The system is extremely complex and the original descriptions cited above are detailed, Consequently, no adequate r6sum6 can be given here and reference must be made to the original papers for detailed comparative information. The system of muscles in the cave-dwelling Spelaeomysis longipes is simplified according to Nath (1974). This is apparently a result of its lethargic life style in that swimming muscles are either absent or atrophied. The muscles of the head of Praunus JIexuosus are described by Mayrat (1955).

Descriptions of the gross morphology of the nerve cord of Gnathophausia longispina and Eucopia australis are given by Sars (1885). Chun (1896) describes the thoracic portion of the cord of Arachnomysis leuckartii. The cerebral ganglia lie anteriorly and dorsally to the mouth and the large paired optic nerves and the smaller paired antennulax and antenna1 nerves originate from its anterior end (Fig.

7.

INTERNAL ANATOMY

123

34). Circumoesophageal commissures connect the cerebral ganglia with the nerve cord. Some of the thoracic ganglia are fused together but the six abdominal ganglia are distinct in Gnathophausia longispina (Fig. 34). Dahl (1956) shows a sagittal section through the cerebral ganglia of Praunus Jlexuosus to demonstrate the positions of the tritocerebrum, deutocerebrum and the anterior and more dorsally sited protocerebrum. The histological structure of the cerebral ganglia of Eucopia and Boreomysis species is described by Hanstrom (1948) and Siewing (1956). The eyes of mysids vary considerably in their degree of development and size relative to the body and gross form. They are reduced to a coalesced plate, dioptric elements usually being absent, in species of the genera Scolamblyops, Michthyops, Pseudomma and Parapseudomma. The eyes exist as separate plates, again with the visual elements reduced or absent, in species of the genera Amblyops Ambtyopsoides, Mysimenzies, Paramblyops, Dactylerythrops and Calyptomma. The eyes of one species of Ceratomysis and Hansenomysis are separate but reduced to spinous processes (Fig. 4.10). Petalophthalmus species have very small eyes (Fig. 4.12) while those of Hansenomysis species are even more reduced and plate-like (Fig. 4.13). Other genera that FIQ.31. Musculature of Gnathophausia zoea (after Daniel, 1933). m, muscle. TT, thoracic transverse m; I , thoracico-abdominal m; 11, anterior oblique rn 1; IIa, external arm of anterior oblique m 1; IIb, oblique transverse m, anterior oblique m 1;IV, anterior oblique m 2; IVa, external arm of anterior oblique m 2; IVb,oblique transverse m, anterior oblique m 2; IVc, posterior auxiliary m, anterior oblique m 2 ; V, central m 1; Vs, dorso-lateral m, abdominal segment 1; VI, transverse m, abdominal segment 1; VIII, central m 2; IX, oblique transverse m 1, anterior oblique m 3; X, anterior oblique m 3; Xa, external arm of anterior oblique m 3 ; Xb, auxiliary m, anterior oblique m 3; Xc, posterior auxiliary m, anterior oblique m 3; XII, dorsolateral m, abdominal segment 2; XIII, transverse m, abdominal segment 2; XIV, posterior oblique m 3; XV, central m 3; XVI, anterior oblique m 4; XVIa, external arm of anterior oblique m 4; XVIb, auxiliary m, anterior oblique m 4; XVIc, posterior auxil'iary m, anterior {obliquem 4; XVII, ,oblique transverse m 2, anterior oblique m 4; XVIII, dorso-lateral m, abdominal segment 3; XIX, transverse m, abdominal segment 3; XX, posterior oblique m 4; XXI, central m 4; XXII, anterior oblique m 5; XXIIa, external arm ,of anterior oblique m 6; XXIIb, auxiliary m, anterior oblique m5; XXIIc, posterior auxiliary m, anterior oblique m 5; XXIII, oblique transverse m 3, anterior oblique m 6; XXIV, dorso-lateral m, abdominal segment 4 ; XXV, transverse m, abdominal segment 4; XXVIa, oblique transverse m 4, anterior oblique m 6; XXMI, central m 5; XXVIII, anterior oblique m 6; XXVIIIa, external ,arm of anterior oblique m 6; XXVIIIb, auxiliary m, anterior oblique m 6; XXVIIIc, posterior auxiliary m, anterior oblique m 6; XXIX, central m 6; XXX, anterior oblique m 7; XXXa, external arm of anterior oblique m 7 ; XXXb, auxiliary m, ,anterior oblique m 7; XXXc, posterior auxiliary m, anterior oblique m 7; XXXIII, posterior auxiliary m, abdominal segment 6; XXXIIIa, flexor muscle of telson; XXXIIIb, longitudinal muscle, abdominal segment 6; XXXIV, auxiliary ,muscles, abdominal \segment 6 ; XXXVIII, dorsal flexor muscles of telson; XXXIX, ventral flexor muscle of telson.

Superficial ventrot musEles

Thwack muscles \ 1-6 \

,'

FIG.32. Musculature of Lop?wga&er t y p i c w . Upper figure showing dorsal, lateral and superficial ventral muscles; lower figure showing main ventral muscles. Nomenclature of muscles is given in the legend to Fig. 31 (after Daniel, 1933).

-

Lateral muscles

sor muscle of tail lobe

flexor muscle of tail lobe

J

FIG.33. Musculature of P r a u n u s j l m w . Upper figure showing main and lateral dorsal and lateral muscles; lower figure showing main ventral muscles (after Daniel, 1928). I, anterior thoracic m; Ia, thoracic longitudinal m 1; Ib, thoracic longitudinal m 2; 11, oblique “transverse” m 1 ;111, anterior oblique m 1 ; IV, thoracic central m ; IVa, thoracic transverse m; V, posterior oblique m 1; VI, central m 1 ; VII, oblique “transverse” m 2; VIII, anterior oblique m 2 ; IX, posterior oblique m 2; X, anterior central m 2; XI, oblique “transverse” m 3; XII, anterior oblique m 3; XIII, posterior central m 2; XIV, posterior oblique m 3; XV, subsidiary m, anterior oblique m 3; XVI, anterior central m 3; XVII, oblique transverse m 4; XVIII, anterior oblique m 4; XIX, posterior central m 3; XX, posterior oblique m 4 ; XXI, subsidiary m, anterior oblique m 4 ; XXII, anterior central m 4; XXIII, oblique transverse m 5; XXIV, anterior oblique m 5; XXV, posterior central m 4; XXVI, posterior oblique n, 5; XXVII. subsidiary m, anterior oblique m 5; XXVIII, anterior central m 5; XXIX, oblique transverse m 6; XXX, anterior oblique m 6; XXXI, posterior central m 5; XXXII, XXXIV, XXXV, flexor muscles of abdominal segment 6; XXXIII, transverse m, abdominal segment 5; XXXVI, subsidiary flexor m, abdominal segment 6; XXXVII, transverse m, abdominal segment 6; XXXVIII, dorsal flexor muscles of telson; XXXIX, ventrsl flexor muscle of telson.

128

THE BIOLOGY O F MYSIDS C

\

9

Fra. 34. The nerve cord of (7nathphazcsia Zongispim (after Sam, 1885). a, f h t abdominal ganglion; c, cerebral ganglia.

have species with markedly reduced eyes that are more or less normal in shape are : Antromysis, Bathymysis, Chalaraspidum, Dactylamblyops, H yperamblyops , Parambl yops , Pseudambl yops , Pseudochalarasp idum, Pseudomysis, Spelaeomysis, Stygiomysis, Teraterythrops and Troglomysis. Zharkova (1970) has studied the histology and reduction of the eyes in Boreomysis californica, B . inermis, Eucopia australis, E . grimaldii, Hyperamblyops nana, Petalophthalmus armiger, Dactylamblyops pellucida and D . tenella. Various stages of reduction are evident ranging from an ommatidium that is complete except that there is no lens, through an ommatidium with no lens and no nerve connection to the rhabdom, t o an ommatidium where both the lens and the rhabdom are absent. Some species have eyes of peculiar shape; those of some species in the genera Erythrops and Hyperythrops are kidney-shaped and red in colour, while the eyes of Heteromysoides species are of the form shown in Fig. 10.26. Divided eyes where the visual elements occur in two quite distinct regions (Figs 8.20, 8.22) occur in species of the genera Anisomysis, Caesaromysis, Carnegiemysis, Echinomysides, Echinomysis, Euchaetomera and Euchaetomeropsis. A few other species also have modified compound eyes that are usually described as having accessory eyes (Fig. 7.12); these are in the genera Dioptromysis, Kainmatomysis and Leptomysis. Casonova (1977) found that the eyes of Eucopia unguiculata (hanseni) are larger, relative t o the body size, in individuals living in the north-eastern Atlantic than those living in the Mediterranean. The number of ommatidia in the eye is related to carapace length of the animal, Atlantic individuals of carapace length 3.5-7-5 mm having 50-220 ommatidia while the Mediterranean individuals of carapace length 3.5-6.5 mm have 50-130 ommatidia. According to Hallberg (1977), there are about 1500 ommatidia in each eye of Erythrops serrata, about 600 in each eye of Mysidopsis gibbosa, some 1600 in the eye of Neomysis integer, about 200 in the eye of Praunusflexuosus and about 5000 in the eye of Siriella norvegica. Thus the numbers of ommatidia present in an eye vary widely between species but the significance of this is not known. The most detailed studies of the general histology of mysid eyes are those of Mayrat (1956a, 1962) on Praunus jlexuosus, Eguchi and

7. INTERNAL

ANATOMY

129

Waterman (1966, 1973) on those of Mysidium gracile and Mysis relicta, Kulakovskii (1969) on 'those of M . oculata and Hallberg (1977) on those of Erythrops serrata, Mysidopsis gibbosa, Neomysis integer, Praunus Jlexuosw and Siriella norvegica. There is little remarkable about the general structure (Fig, 35) because it is comparable to that of the compound eyes of other crustrtceans. Hallberg (19771, in his comparative study of five species, found that the ommatidia of the different species are constructed on the same general plan although they vary considerably in detail. Variation exists in the number of retinular cells, the presence or absence of an epirhabdom and in the arrangement, and distribution of &hepigment. The detailed structure of the optic neuropiles (lamina ganglionaris, medulla externa and medulla interna) of Lophogaster longirostris, Mysis oculata, M . relicta, M . litoralis, Neomysis integer, Praunus Jlexuosus and Erythrops servatus (serrata) was examined by Elofsson and Dahl (1970). These ganglia completely fill the eyestalk (Fig. 35) and conform to the general malacostracan pattern. In reduced eyes, such as those studied by Zharkova (1970), there is a tendency for the medulla terminalis and medulla interna t o be closer to the cerebral ganglia. The reduced eyes of Amblyops abbreviata, Boreomysis scyphops and Pseudomma afine were examined histologically by Elofsson and Hallberg (1977). They show that this reduction is not a negative response to an aphotic environment but that the eyes are adapted to reception of low intensities of non-polarized light. The eye of Boreomysis scyphops is dish-shaped (Fig. 35), with an enlarged retinular layer consisting of some 3000 ommatidia. The optic neuropiles are well developed although small relative to the size of the retinular.layer. The ommatidia lack a dioptric apparatus and are composed primarily of retinular cells forming 8 cylinder with microvilli protruding into the central chamber. The axons from the ornmatidia pass through a layer of red pigment cells, which serve as a basement membrane, to the lamina ganglionaris. The eye of Amblyops abbreviata (Fig. 35) is different, contains some 1000 ommatidia and also has a well-developed retinular layer, pigment cell layer and optic neuropiles. The eye of Pseudomma afine is of similar construction to that of Amblyops abbreviata but has no pigment cell layer adjacent to the retinular layer. The enlargement of the photosensitive areas by the development of microvilli enhances the reception of low levels of light intensity. Consequently, Elofsson and Hallberg argue that these eyes are positively adapted to the low light regimes of the deep environments. The retinal pigments (Fig. 35) allow dark adaptations of the eyes, as demonstrated by Beeton (1959) in Mysis relicta. Nicol (1959) tested

D

FIQ.35. Compound eyes. P r a u n w $ e m o s w : A, schematic section of an ommatidium and B, of the eye. Boreomysis scyphops: C, horizontal section of the eye. Amblyops abbreviata: D, sagittal section of the eye. (A and B after Mayrat, 195th; C and D after Elofsson and Hallberg, 1977). bm, basement membrane; bv, blood vessel; oc, corneagenous cells and cone cells; cor, cornea; cr cn, crystalline cone, dp; distal reflecting pigment cell; dpc, distal screening pigment; lg, luminu ganglionurb; m, muscle; me, medullu externu; mi, medullu i n t e r n ; mia, accessory medulla i n t e r n ; mt, medulla terminalis; oa, optic artery; p, pigment layer; pc, proximal pigment, r, retinular layer; rh, rhabdom; sg, sinus gland; sp, sensory pore; X, sensory pore X-organ (organ of Bellonci).

7.

INTERNAL ANATOMY

131

Praunus neglectzls and found that the minimum intensity of light to which it responded was 29 x 10" pW/cm2 receptor surface a t 10 cm; the colour of the light was blue, the filter used having a spectral range of 420-540 nm with a maximal emissioh at 475 nm. The reaction time of Mysis relicta t o light of various wavelengths was tested by Beeton (1959) who found that they reacted most quickly to wavelengths close to 515 nm and 395 nm. He suggests that this species probably has two visual pigments. Herman (1962) investigated the spectral sensitivity of the eyes of Nwmysis americana over a narrower spectral range that did not include wavelengths as short as 400 nm. He fhund highest sensitivity at a wavelength of 515 nm. Mysids do not have a nauplius eye nor a dorsal frontal organ (Elofsson, 1965). They do, however, have paired ventral frontal organs situated within the bec ocellaire, which is located a t the anterior end of the body between the bases of the eyestalks and antennules (Fig. 36). Elofsson examined this structure in Lophogaster longirostris, Eucopia sp., Boreomysis arctica, Arcitaeomysis sp., Mysis litoralis, M .

FIG.36. Sagittal section through the anterior part of the head of B o r w m y A a r c t h to show bec owllaire (after Elofsson, 1965). a, antennular nerve; b, cerebral ganglia; c, cells of ventral frontal organ; om, compressor muscles; r, rostrum; vf, ventral frontal organ.

132

THE BIOLOGY OF MYSIDS

oculata, Schistomysis ornata, Praunus Jlexuosus and Neomysis integer. The organ has a similar structure in all species with cells along its length except in Boreomysis arctica (Fig. 36) where the cells are restricted to the distal region. The sensory pore X-organ (SPX-organ) and sinus gland occur within the eyestalk (Fig. 35), the SPX-organ being located towards the dorsal side of the eye, the sinus gland towards the ventral side. Mayrat (1956a), who examined these organs in Praunw Jlexuosw, reviews the earlier literature. Kulakovskii (1969, 1971, 1978) describes their occurrence in the eyestalk of Mysis oculata and their hormonal control over the chromatophores. The SPX-organ of Boreomysis arctica consists of a group of cells and a vesicle, surrounded by a connective tissue sheath, located near the base of the sensory papilla of the eyestalk. The organ is connected by afferent nerves with the medulla terminalis. Neurosecretions produced by the cells of the organ are transported to the vesicle where release is probably effected by the afferent nerves. Dahl and Mecklenburg (1969) suggest that this organ in mysids should be considered not only as a neurosecretory organ but also as a neurohaemal organ. The SPX-organ of Mysis relicta, connected with the medulla terminalis, has “onion bodies” present (Hogstad, 1969) although these were not found to be present within the organ of Boreomysis arctica (Dahl and Mecklenburg, 1969). Kulakovskii (1969) shows an X-organ in the medulla terminalis (MTGXorgan) of Mysis oculata and Gabe (1966) found this ganglionic organ in Siriella sp., Praunus Jlexuosus and Paramysis sp. Hogstad (1969) did not observe it in the eye of Mysis relicta; there was, however, a group of cells situated medio-laterally between the medulla externa and medulla interna, that were assumed to be the medulla interna ganglionic X-organ (MIGX-organ). The sinus gland, described by Hogstad as biscuit-shaped, lies in the lateral-ventral region of the eyestalk close to the medulla interna and medutla terminalis (Fig. 35). Bioluminescence has been reported in several mysids. Illig ( 1905) describes a bioluminescent gland in the maxilla of Gnathophausia ingens. Glandular cells secrete material into a reservoir, the walls of which have muscles attached to them. The reservoir opens to the exterior near the base of the exopod of the maxilla. Tett and Kelly (1973), quoting Harvey (1952), state that, in addition to Gnathophausia species, species in the genus Gastrosaccus and possibly Siriella are luminescent. No modern information is available and investigations of the occurrence of luminescent organs in the Mysidacea are required. Another pair of glands, of a very different nature, the excretory or antenna1 glands are present in the basal segments of the antennae.

7.

133

INTERNAL ANATOMY

They are of the form found in other crustaceans, consisting of an end sac connected by a urinary duct to a terminal vesicle that opens at a papilla to the outside of the animal (Fig. 37). The glands of Lophogaster typicus and Hemimysis Eamornae are described by Cannon and Manton ( 1927b), those of Gastrosaccus spinifer, Mysis relicta, Praunus jlexuosus, P. inermis and Mesopodopsis slabberi by Vogt (1932, 1933) and those of Eucopia species by Siewing (1956). The end sac is oval in shape and has an internal network of blood lacunae. It connects with an irregularly shaped urinary duct, short in Mysis relicta but notably longer in Mesopodopsis slabberi.

P

t

es

m

es

m

FIQ.37. The excretory organs (antenna1glands) of left, Guatrosaccus spinifer and right, Myaie relicta (after Vogt, 1932, 1933). es, end sac; m, mandible;p, external opening; t, terminal vesicle; ud, urinary duct.

A gland, termed a Y-organ, is described in the ventral region of the head of Praunus jlexuosus, Paramysis sp. and Siriella sp. by Gabe (1952, 1956). This gland appears to be analagous with the ventral gland of insects and secretes a hormone controlling the moulting cycle. There is some evidence suggesting that mysids are able to locate, by chemosensory mechanisms, sources of food. The presence of excretory material, detected by similar sensors or sensilla, may also convey useful information to the mysid. Clutter (1969) hypothesized the presence of chemo- and mechanoreceptors in mysids when examining the swarming behaviour of Metamysidopsis elongata. Fuzessery and Childress (1975) investigated the amino acid feeding response thresholds of Gnatlwphausia ingens and some other crustaceans. Concentrations of amino acids (L-glutamic acid, taurine and DL alpha aminon-butyric acid) of 10-10-10-11 M initiated feeding behaviour. The dactyl receptors of G, ingens had a response threshold at 6 x 10-8 M, while antenna1 receptors had a threshold a t 5 x lo-' M. Fuzessery and Childress discuss the implications of these observations including the

134

THE BIOLOGY OF MYSIDS

possibility that this species, and other species, can orientate to a chemical gradient in order to locate food. No description of the chemosensors involved in the detection of the amino acids is given by Fuzessery and Childress. The existence of sensilla on the antennae, mouthparts and thoracic legs of crustaceans has been known for some time. Crouau (1978) and Guse (1978) describe the histological structure and nervous innervation of hair sensilla on the antennae of Neomysis integer. The double and triple innervated hairs are probably mechanoreceptors, while the hairs with many sensory cells are probably mechanochemoreceptors or chemoreceptors. There are, however, also many sensilla and openings of sub-integumental glands distributed over the whole of the integument of the body of crustaceans (Mauchline, 1977s; Mauchline and Nemoto, 1977; Mauchline et al., 1977). These often have distinctive patterns of distribution that are peculiar to families, genera and, in some cases, species. The distribution of such organs in the integuments of Eucopia sculpticauda and Praunus jlexuosus is shown in Fig. 38. The integuments have been digested in hot potassium hydroxide which removes all the soft tissues. They are then cut along the ventral midline and the carapace and dorsal and lateral (tergal and pleural) regions of the abdominal segments laid flat, outer surfaces upwards, on microscope slides for examination. The tissues of the glands and sensilla are digested by the hot potassium hydroxide and only a hole (pore) is left at the site in the integuments where the sensilla or gland was present. The hole or pore is only an indication of a connection between the sub-integumental tissue and the outside environment. This connection can be one of the following: a tubular duct of a subintegumental gland, that releases its secretions to the outside environment ; a tube connecting a sensillum with its axons and sensory neurons in the hypodermis; a tube through which nerve fibres from the peripheral nervous system pass outwards to that part of the sensillum that is external to the integument. These pores occur over the whole of the integument, including the exposed tergal areas of the posterior thoracic segments (Fig. 38). There is a varying degree of bilateral symmetry, most evident in the distribution of conspicuously large pores or in that of conspicuous groups of pores. The distributions of conspicuous pores or groups of pores in species in different genera are shown in Fig. 39. Eucopia sculpticauda and Praunus jlexuosw are included. The distributions show considerable variations between genera. Dr M. Murano supplied 11 Siriella spp., 7 Neomysis spp. and 10 Acanthomysis spp., and the distributions of the pores were examined. The distributions shown in

7.

INTERNAL ANATOMY

135

FIG.38. Distribution of pores in the treated integument of (left) Praunwflexuow and (right) Eucopia aculpticauda (Mauohline, 1977).

Fig. 39 contain pores common to all species examined within each of these genera. There was remarkably little variation in the general pattern between species within these genera and this is emphasized by the identical overall patterns shown for the closely related genera Neomysis and Acanthomysis (Fig. 39). Detailed examination of the distribution apd numbers of pores in species within these genera did produce specific differences in some cases, but not in others. The two species of Schistomysis examined (Fig. 39) are rather different and so larger specific differences may occur in some other genera.

... .... -..

-.. ....

.%

& ,-.

%.

C B

!2

*...

H.

............ PT

................. ...........

9.

...........

*.

.r

...........

.__ -- .-..

-.-...... c

1

...-....

2

..-

.v.

._ ..-* _-

I;,

3 4

5

6 Ery f hrops serrafa

Eucopia sculpficauda

Schisfomysis spiritus

.-. ....

9

-

....-_.

S. ornufa r

9

....-..-.

-... ..- ./ . I

#

Praunus f lexuosus

Kaf ery fhrops oceanae

"c--

........... ..

..."-....

..,,......*.. -. ...... :,.... ......

A canthornysis soecies

Neomysls soecies

.-.-.

............

.-..--.

:.1

*.

Mysidopsis gibbasa

. I . , .

.......... /

......-.. ..

...........

............

..........

.........

..........

..........

5.

1 FIQ.39. Distribution of groups of pores in the carapace (C), posterior thoracic segment (PT) and the six abdominal segments 1-8.

7.

INTERNAL ANATOMY

137

The sub-cuticular and intra-cuticular structures of the sensilla and glands that were present a t the sites of these pores before digestion of the integument by potassium hydroxide are unknown. The surface of the fresh integument has been examined and a variety of sensilla and gland openings recognized. There are feather-like setae that are probably hydrodynamic receptors. They tend t o occur in dorsal anterior regions of the carapace and other dorsal regions of the body, including the abdomen; Debaisieux (1947) draws some on the dorsal surface of the endopod of the uropod and shows nerves travelling to them (Fig. 40ds). The rows of pores occurring across the dorsal region of the carapace and abdominal segments in many species (Fig. 39) often represent the sites of small conical structures with or without a pore a t the apex; these are thought to be chemosensory in function. Much further work is required, especially on the histological structure of these organs. It seems probable, however, that the mysids are monitoring various parameters of their three-dimensional environment, through the use of integumentory receptors. Clutter (1969) suggests that Metamysidopsis elongata, in common with other crustaceans, uses chemical cues for mgting. The newly moulted female appears to be located along a chemical gradient by the male. The specific receptors involved have not been identified. One further pair of large sense organs are present in the Sub-Order Mysida except in the family Pekalophthalmidae, but are absent in the other Sub-order, Lophogastrida. They are the statocysts, or gravity receptors, present in the endopods of the uropods (Fig. 40). The histology of the statocyst of Praunus jlexuosus is described by Debaisieux (1947, 1949) who reviews the earlier literature. The statocyst comprises a vesicle inside which is a lith suspended on sensory hairs. The lith is secreted by the mysid a t the moult that terminates Stage I11 of the larval development ; this moult takes place shortly after emergence from the marsupium. The lith consists of a protein matrix surrounded by a calcareous shell that is rich in calcium fluoride (Enbysk and Linger, 1966) and has an undetermined organic base. The shell is penetrated by the 50-60 sensory hairs by which the lith is suspended in the vesicle. These hairs originate from a ventral sensory cushion (Fig. 40) and are inserted along the outside lateral edge, relative t o the animal's body, of the lith (Fig. 40). The planes of the sensory cushion and lith in the two statocysts are inclined towards each other, ventromedially, so that they are a t an angle of 50-55" t o the horizontal longitudinal plane of the animal. Consequently, the sensory hairs are under tension when the animal is orientated horizontally. Neil (1975~) discusses the shearing forces acting on the sensory hairs during

138

THE BIOLOGY OF MYSIDS

A

\

C

\ \ \

,'550',

FIQ.40. The statocyst of Praunus ~ V e x w a w r .A, uropods showing position of stetocysts in the endopods. B, sagittal section through the lith within its vesicle. C, diegrammetic transverse sections of the endopods to show the orientation of the litha relative to each other and to the horizontal. (A end B after Debaisieux, 1947; C, after Neil, 1975~).c, sensory cushion; ds, dorsal sensory aetae; 1, lith; m, muscle; n, nerve; sh, sensory hairs of lith; v, vesicle.

rotation and tilting of the animal and shows that the neural control system sunimates the afferent signals from both statocysts. The statocysts operate in conjunction with the eyes to control orientation of the mysid in space (Neil, 1975a-c). The shearing stimuli over the sensory cushion in each of the two statocysts result in opposite turning reactions because of the mirror inclinations of the cushions. This bi-directional input to the statocyst control system, however, results in an essentially uni-directional output to the eye stalks. Applying visual stimuli, such as a vertical light beam, modifies the eye movements but leaves their general pattern unaltered, and the orientation of the animal is normal. Orientation was altered in Mysidium gracile by applying a horizontal light beam; the mysids compromised and orientated a t about$45" to the horizontal (Jander, 1962). Rice (1961, 1964) found that the effects of gravity dominated those of light in the orientation process. Increase in hydrostatic pressure caused the mysids to move upwards in the water column, even when illuminated from below. This reaction took place in Leptomysis lingvura, Schistomysis spiritus, Praunus jiexuosus, P . neglectus and to a, considerable

7. INTERN&

ANATOMY

139

but lesser extent in SIiriella urmatu. An additional sensory input is probably used for purposes of orientation by, for example, epibenthic species ; this input is from tactile receptors on the dactyls of the thoracic legs (Foxon, 1940). Consequently, orientational responses probably result from integration of inputs not only from the statocysts and eyes, but also in certain circumstances, from additional inputs from integumental receptors, especially tactile receptors on the thoracic legs.

CHAPTER 8

PHYSIOLOGY A N D RESPONSES TO PHYSICAL-CHEMICAL PARAMETERS OF THE ENVlRONMENT Mysids exhibit a diversity of behavioural patterns. Their behavioural responses are elicited by two general classes of environmental stimuli; the first consists of changes in the physical and chemical parameters, while the second consists of stimuli originating from other organisms, including other mysids. In addition, of course, the internal physiological state of the individual also has an influence on its behaviour a t any one instant of time. Consequently, the physiology of the animals and their responses to the physical-chemical parameters of the environment will be discussed in this chapter. This is followed by an examination of more general aspects of the behaviour of individuals and populations in the next chapter. Swimming is effected by beating the exopodites of the thoracic legs. Many female mysids have rudimentary pleopods while in many males the pleopods, if developed, are modified as secondary sexual characters, and are not used in swimming. The beating of the thoracic endopodites generates the respiratory currents as well as providing propulsion for the body. Laverack et al. (1977) have shown that the beat of an exopodite has a modified rowing action. It consists of a power stroke, a return stroke, and a pause. The beat frequency in Praunus jlexuosus is about 4 Hz. The co-ordination between successive exopodites on the same side of the thorax and between the exopodites of corresponding legs on opposite sides of the body is strong. The speed of swimming of mysids has not been measured in many species. Metamysidopsis elongata of 4-7 mm total body length (a length representitive of many mysids, see Fig. 20, p. 72) can swim at speeds up to 13cm/s but normal cruising speed is probably within the range of 5-6 cm/s (Clutter, 1969). Acanthomysis species of 5-9 mm total length can maintain speeds of up to 20 cm/s in the laboratory, although Clutter considers that they normally swim at speeds of 5-7 cm/s in the sea. Another species of comparable body size, Mysidium columbiae, can swim at speeds of

8.

PHYSIOLOGY AND RESPONSES TO THE ENVIRONMENT

141

up to 15 cmjs (Steven, 1961). Much slower swimming speeds are mentioned by Foulds and Roff (1976) in Mysis relicta of 10 mm total body length; they measured respiration rates a t swimming speeds in the ranges 14-1.6 and 2.1-3.6 cm/s. Robertson et al. (1968) estimated a swimming speed of 5-10cm/s in this species while observing its behaviour from a submarine. Consequently, the maximum swimming speed of this species remains unknown. Paramysis intermedia could not maintain its position in water currents exceeding velocities of 40 cm/s (Boroditch and Havlena, 1973). Species and individuals vary in the amount of time they spend swimming. All species are slightly denser than the surrounding seawater; Morris (1972) found that the density of Eucopia sculpticauda was 1.024. An epibenthic habit influences the amount of time spent swimming and low light intensity has been shown to have an inhibitory effect on swimming. Some species appear to swim almost continuously while others have what are probably endogenous rhythms of activity. For instance, Hobson and Chess (1976) found that Siriella pacijica is a nocturnal carnivore that lives on the surface of the seabed among the kelp, Macrocystis pyrifera, during the day, but forages outside the kelp beds at night. A co-occurring species Acanthomysis sculpta, however, fed by day and night usually entirely within the confines of the kelp beds and showed no periods of increased or decreased activity. Burrowing species of Gastrosaccus rest during the day in the sediment, but emerge at night to swim; this is an endogenous rhythm of nocturnal emergence and Macquart-Moulin ( 1 9 7 7 ~ ) found that this pattern of swimming activity persists under experimental conditions of constant darkness for 10 days in G. mediterraneus and for about 50 days in G. spinifera. Most hyperbenthic species probably rest for periods on the surface of the sediment or local algae. Species such as Praunus$exuosus can be observed doing this in shallow water during the day for one to several hours a t a time. Two hyperbenthic species, Leptomysis gracilis and Siriella jaltensis, were found to have no endogenous rhythm of activity and a further species, S . clausii, only had a weak rhythm (Macquart-Moulin, 1973b). Anchiah a agilis, which according to Macquart-Moulin, is hyperbenthic in the Mediterranean but which is more strictly pelagic in British waters, also exhibited no endogenous rhythm of swimming activity. Consequently, it is probable that endogenous rhythms are confined to hyperbenthic species living among the macro-algae in shallow habitats and other species living in specialized situations, such as burrows. Hyperbenthic species living in deeper habitats without sheltered regions such as those provided by macro-algal beds, probably have

142

THE BIOLOGY OF MYSIDS

irregular periods of rest on the sediment surface. Pelagic species probably swim more or less ‘continuously. The levels of energy expended in swimming have been measured in two mysids. Klyashtorin and Kuz’micheva (1975) found that the coastal Neomysis mirabilis expended energy during swimming a t a level equivalent to 1.3 times that of the rate of standard metabolism. They conclude that planktonic crustaceans weighing 1-100 mg in body weight and swimming a t rates characteristic of periods of prolonged continuous swimming have total energy expenditures that exceed levels of energy consumption at relative rest by factors of 1.5-2.0. Comparable factors were determined for the large bathypelagic species Gnathophausia ingens by Quetin et al. (1978). They found a factor of 2.3 a t a rate of swimming activity that the animal could maintain continuously for a period of a t least 14 days. This factor increased to 2.9 a t the maximum rate of swimming activity observed in this species. Some 3% of its metabolic scope was required by G . ingens to maintain its position in the water column and about 40% for other swimming activity. Quetin et al. concluded, contrary to their previous convictions, that G. ingens swims more or less continuously a t a steady and relatively rapid rate and does not have a lethargic life style. Foulds and Roff (1976) also examined levels of energy expenditure during swimming. They, however, used the rate of routine metabolism as the reference rate. They defined routine metabolism as present when “a behaviour pattern of resting interspersed with spontaneous activity under conditions of minimal external stimulation” existed. A maximum increase of 1.1 times the routine rate waa present in the freshwater Mysis relicta when swimming a t speeds of 1-0-1-6 cm/s and 1.2 times the routine rate a t swimming speeds in the range 1.2-3.6 cm/s. These swimming speeds are probably low relative to the maximum speeds of this species. Rhythms of activity may be interrupted temporarily by moulting although there is no information on this aspect in mysids. Nouvel (1957b) discusses the process of moulting in a range of species and Saudray (1972) examines the effect of moulting on the functioning of the mandibles of Praunus jlexuosus. Holmquist ( 1959b) considered that Mysis relicta maintained in aquaria at 4°C had intermoult periods ranging from 17 to 24 days and sometimes from 28 to 30 days. Blegvad (1922), rearing individual Praunus inermis and P . Jtexwsus in aquaria, observed intermoult periods of irregular duration; his data are not presented in a form that will allow more detailed exatmination. The intermoult periods appear to range from 15 to about 60 days, a modal range being approximately 20-30 days. The duration of the

8. PHYSIOLOGY AND RESPONSES

TO THE ENVIRONMENT

143

intermoult period was found to increase logarithmically when plotted against body length or successive moult number in euphausiids and other decapod crustaceans (Mauchline, 1977b, c). The intermoult periods determined in Praunus JEexuosusby Nouvel and Nouvel (1939) and Metamysidopsis elongata by Clutter and Theilacker (1971) are shown in Fig. 41 in relation to total body length. Dr R. Gaudy kindly

30r 1-003

10

Carapace length ( rnm) 20 30 40

50

Gnathophausis ingens

5 5°C 6 5°C 7 5OC

20 -200

10-100 P L a Q ,

4-

Praunus f/exuosus

-

0 Mefarnysidopsis elongofa

A

Melongofa

2 ' 1 2

I

4

'

I

6

I

I

8

I

"

10

]

12

I

14

1

I

16

I

Total length (rnrn)

FIQ.41. The durations of the intermoult period related t o body size of three species of mysids. The durations of the periods of fhzathophausia ingens are 100-250 days and are related to carapace length and determined a t 3 temperatures. The regression, equations of the lines are given in Table XXIII. Data on Praunw fZexuosus from Nouvel and Nouvel (1939), on Mebmyeidopsis elongab from Clutter and Theilacker (1971) and on G m t h p h a w i a ingem from Childress and Price (1978).

allowed me to examine his data (Gaudy and Guerin, 1979) prior to publication ; the relationship between the duration of the intermoult period and successive moult number a t several temperatures is shown in Fig. 42. The regression equations for these relationships are given in Table XXIII, the regression coefficients all being significant a t the 1.0-0-1% level. Childress and Price (1978) have determined the intermoult periods of the bathypelagic Gnathophauaia ingens. These ranged

144

THE BIOLOGY OF MYSIDS

30 r

Y 20 -

-"z 10-

0

-

-

0

u

0 .-

b a

c

-

7-

-

3

0

E

4-

0)

c

C -

Lepfomysis hnqvuro 10°C A 14°C o 18°C A 22°C

214°C 018°C A 22°C A

A

1 -

1

1 2

1

1 4

1

1 6

1

1 8

1

1

1

1 10 Moult number

3

5

7

9

FIG.42. Duration of the intermoult period related to successive moult number at various experimental temperatures in Hemimysis spetuncola and Leptotmysis lingvura. Data from: Gaudy and Guerin (1979).

from about 105 to 210 days, dependent upon the sizes of the individuals and the experimental temperatures. They expressed the relationships at the three temperatures as:

+ 0.00051L3 6.5"C D = 137.7 + 0*00050L3 7 ~ 5 ° C D = 120.5 + 0*00038L3 5.5"C

D

= 166.3

where D is the duration of the intermoult period in days and L3 is carapace length to the third power. These equations have been transformed in Table XXIII for comparison with those of the other species. The most notable feature of the equations for this relationship in G. ingens is that their slope ( b ) is markedly reduced, not only relative to that of the other mysids in Table XXIII but also relative to other

TABLEXXIII. REGRESSION ANALYSES OF THE LOGARITHM OF THE ~ ~ R M PERIOD O ~ T (IN DAYS)ON BODYLENUTEOR SUCCESS~VE MOULTNUMBER Regression constants logy = bx f a Species

Independent measurement

Gnuthophauaia ingens a t 5.5"C a t 6.5OC at 76°C

CL CL CL

Hemimysis speluncoh at 10°C at 14°C at 18OC a t 22°C

MN

m

MN MN

M~ASWMENT OF

a

b

2.1700 2.0804 2-0289

0.0040

10 10 9 10

0.5575 0.3517 0.1602 0.1 112

0.0831 0.0870 0.0964 0.0907

0.9169* * * 0*9249*** 0*9693*** 0.9315***

Gaudy and Gaudy and Gaudy and Gaudy and

Guerin (1979) Guerin (1979) Guerin (1979) Guerin (1979)

5 7

0.6975 0.4239 0-3036 0.0117

0.1305 0.0780 0.0661 0.0873

0.9874** 0*8090*** 0.8795*** 0.9359***

Gaudy and Gaudy and Gaudy and Gaudy and

Guerin Guerin Guerin Guerin

0.3604 0.3886

0.0699 0.0850 0.0676

0.9795** * 0.5987*** 0.9717* * *

Clutter and Theilacker (1971) Clutter and Theilacker (1971) Nouvel and Nouvel (1939)

n

Leptomysis lingvura a t 10°C a t 14°C at 18°C at 22°C

MN MN MN

10

Metamydopsis elon.gata females males Praunua jlexuo;lu8

TL TL TL

5 6 11

MN

Value and signijkance of

A

7

0.0048

Source of data

Childress and Price (1978) Childress and Price (1978)Childress and Price (1978)

0.0035 0.0035

(1979) (1979) (1979) (1979)

TL, total length; CL, carapace length; MN. moult number; n, number of observations. The value and significance of the regression coefficient t is given.

146

THE BIOLOGY OF MYSIDS

crustaceans (Mauchline, 197713, c). The relationships in adult euphausiids have values of b,ranging from 0.0058 t o 0-147, in decapods other than panulirid from 0.010 to 0.054 and in panulirids from 0.004 to 0.009. Hartnoll (1978) suggests that b decreases in larger decapods and this may also be true in mysids. Conversely, it may be related in some way to the conditions of the bathypelagic environment in the case of G. ingens. Further discussion of this aspect and its effect on growth rates is presented in Chapter 10. The rates of oxygen consumption of mysids during respiration are comparable to those of other crustaceans (Table XXIV). The TABLEXXIV. OXYGEN

Species Gnathophausia ingens Gnathophausia zoea Gnathophawia gigas Gnathophawia gracilis Boreomysis californica Archaeomysis grebnitzkii" Gastrosaccua simulansa Hemimysis lamornae Holmsiella anomla" Hypererythrops sp. Metamysidopeis elongata" Mysis relictaa

Temperature ("C)

pl OJmg wet wt/h

5.5 5.5 5.0 5.5 4.0

0'024-0'48 0.035 0.031 0.027 0.032 0.022 0.033 0.17 0.27 0.49 1.54-2.40 1.8 3.6 0.3 0.79 0.5-0.7

4.0

5.5 5 10 15 27.5 10 20 13 17 13.8 5 4

Neomysis americana Neornysis awatschensisa Neomysis in,teger

a

CONSUMPTION OF

5 5 10 15 10 18 f 0.5

MYSIDS

Reference

Childress (1971a) Childress (1975) Quetin et al. (1978) Childress (1975) Childress (1975) Childress (1975) Childress (1975) Jawed (1973) Jawed (1973) Jawed (1973) Gauld and Raju (1975) Grainger (1956) Grainger (1956) Ikeda (1977) Ikeda (1970) Clutter and Theilacker (1971) Lasenby and Langford 0.4 (1972) Foulds and Roff (1976) 0.21 0.26-0.38 Raymont end Conover (1961) Jawed (1973) 0.21 Jawed (1973) 0.29 Jawed (1973) 0.35 0.15-0.46 Raymont et al. (1966) 0.18- 1.29 Raymont and Krishnaswamy (1968)

Original data on dry weight basis; dry weight equivalent to 20% wet weight.

8.

PHYSIOLOGY AND RESPONSES TO THE ENVIRONMENT

147

bathypelagic species of the genera Gnathophausia and Boreomysis have lower weight specific respiration rates as discussed by Childress (1971b, 1975). Their increased body size relative to that of coastal and epipelagic species does not account for this decrease (Fig. 43) as pointed out by Childress. The haemolymph, which probably contains a haemocyanin, of Gnathophausia ingens has an unusually high affinity for oxygen. Free1 (1978) found that the partial pressure of oxygen a t half saturation (P50)is 1.5 k 0.19 mmHg, and suggests that the P,, may be well below 1 mmHg when this species is living in its normal habitat a t temperatures of 5°C and less. The rates measured in Gastrosaccus simulans by Gauld and Raju (1975) are exceptionally high, but may be partly accounted for by the high environmental temperature, 27.5"C in which this species lives. Chin (1976) obtained rates of respiration in Neomysis awatschensis two or more times higher than those determined by Jawed (1973) in Table XXIV; the conditions of salinity in the experiments differed. Comparable rates of respiration occurred in Paracanthomysis hispida (Lee and Chin, 1976). Klyashtorin and Musayeva (1977) compare estimations of respiration rates of Neomysis mirabilis using an oxygen electrode and the standard Winkler chemical method. Lower rates, lower by 8%, were measured with the electrode but they conclude that its use makes such measurements easier and faster to carry out. Various factors, in addition to swimming activity, affect the respiration rate. The most important is the environmental temperature, increasing temperature being associated with higher respiration rates (Grainger, 1956). Clutter and Theilacker (1971) estimate a Qlo for Neomysis americana of 1.6 and for N . integer and Hemimysis lamornae of 1.9; this latter factor was calculated from the data of Grainger (1956) on H . lamornae. A Qlo of 2.9 for Archaeomysis grebnitzkii and of 1.7 for Neomysis awatschensis are present in Table XXIV. Chin (1971) found the Qlo for Neomysis awatschensis, in experiments where individuals were acclimatized to 5" intervals of temperature in the range 5-25"C, to be between 1-56 and 2.18 over the 5-15"C range and to be 1.17-2.64 over the 15-25°C range. The effect of thermal acclimatization on respiration rates of this species was investigated further by Chin (1974). A Qlo of approximately 2.0 was found by Lasenby and Langford (1972) for the freshwater Mysis relicta. The effects of varying combinations of temperature and salinity or of varying salinity independent of temperature have been investigated by Chin (1972, 1974), Jawed (1973), Foul& and*Roff (1976) and Vlasblom and Elgershuizen (1977). There are optimal combinations of temperature-salinity for each species, as would be expected.

3-00 @

1-00

i $IL

3 c 0 \

3

c

5

- 1

5 010

0 N

F

0.011

n

Arcboeomysis grebnifzkii Gaslrosaccus sirnulans Hemirnysis farnornae Hypererytbrops sp. Metamysidopsis elongafa Mysis relicta 0~ y s i reficta s 0 Neomysis arnericona Neomysis owotscbensis Boreomysis californico

sr

I

I I l l l l 1l0

I

I

I

I I 1 1 1 1

102 Body wet weight ( mg )

t

I

I

G.ingens

I

H

G.qiqas Ggracilis

I

* I

Gnotbopbausia species

I

I

I

I I I I l l

1 o3

I

i

I

I I I I Ill 104

FIG.43. Respiratory rates of mysids related to body wet weight. The two regreasion &es in the left upper part ofthe figure are for Metamy&pia e h g a t a and Mysis d i c t a respectively. The respiratory rat0s of Gnathopha& species are presented in relation to their range of body wet weight. Data from: Grainger, 1956; Raymont and conover, 1961; Ikeda, 1970; Clutter and Theilacker, 1971; LasenbyandLangford, 1972; Jawed, 1973; Childress, 1975; Gauld and Raju, 1975; Foul& and Roff, 1976.

8.

PHYSIOLOOY AND RESPONSES TO THE ENVIRONMENT

149

The different species so far investigated show varying aptitudes for acclimatization to 'extreme values. The degree of oxygen saturation of the water also affects the ra-tes of respiration. Species such as Holmsiella anornula examined by Ikeda (1977) show a high tolerance to oxygen concentrations as low as 10% of saturation. Archaeomysis grebnitzkii and Neomysis awatschensis, two coastal species, were less tolerant, dying at levels of less than 13% saturation (Jawed, 1973). The most tolerant species, with aerobic regulatory abilities, are the bathypelagic GnathophaMia species and Boreomysis calijomica investigated by Childress (1968, 1971a, 1975). Their respiratory rates did not decline markedly until the oxygen concentrations reached the low levels of about 6% saturation. These species inhabit the oxygen minimum layers of the eastern Pacific where oxygen is a t minimal concentrations of about 0.20 ml 0,/1 (equivalent to about 4% saturation). Gradients of pH are associated with these regions of oxygen minimum layers, the pH ranging from 8.3 at the surface to 7.5 or less in the oxygen minimum layers. Mickel and Childress (1978, however, found that changing pH of the water had no effect on the respiratory processes of Gnathphausia ingens. The only activity affected by changing pH was the extraction of oxygen from the respiratory stream; extraction rates were 10-30% higher a t pH 7.1 than at pH of 7.9 and 8-7 but why this was so is unknown. A seasonal change in the respiratory rate of Neomysis integer was demonstrated by Raymont et al. (1966). Minimal levels occurred in September-October followed by successively increasing levels to a pronounced maximum in March. No other information on this aspect is available. The weight specific respiration rates decrease within a species with increasing body size (Chin, 1971, 1974; Clutter and Theilacker, 1971; Jawed, 1973; Lee and Chin, 1976; Vlasblom and Elgershuizen, 1977) and a similar trend is present between species (Fig. 43) as suggested by Ikeda (1970). Shushkina (1973) determined the relationships between respiration and body weight in Neomysis mirabilis in the sea of Japan as:

R

= 0.14

A detailed study has been made of the respiration rate of N . mercedis by Simmons and Knight (1975). They derived a general equation relating its respiration rate (yl O,/animal/h) to changing values of body dry weight (mg), temperature ("C), sex (see below), salinity (% sea water) and month of the year (1-12). Sex was scored 1-5; 1 being for juveniles, 2 for males, 3 for immature females, 4 for non-ovigerous

150

THE BIOLOGY OF MYSIDS

mature females, and 5 for ovigerous females. The equation is:

R

=

2.552+1.779 WT+0-443 T-0-888s-O.Ol8SAL-0.178

M

where R is respiration, W T is dry weight, T is temperature, S is the score for sex, S A L is percentage sea water, and M is the month of the year. They examined the influence of single variables upon the others and upon the respiration rate. The relationship of respiration rate to temperature, salinity and body weight is not constant but varies with season. The simple relationship between respiration and body weight also changes seasonally. They examined all permutations and came to the following general conclusions. The change in the respiration rate in response to changes in either the salinity or temperature is dependent upon the body dry weight of the mysid and its sex. In turn, the influence of body weight and sex depends upon seasonal effects of reproduction and food supply. Changes of temperature a t low salinity and of salinity at low temperature produce larger responses in the respiration rate than the corresponding changes at high salinity and high temperature. An indirect method of measuring respiratory rates is to study the activity of the electron transport system (ETS). Exploratory investigations on other crustaceans have suggested a close correspondence between ETS activity and oxygen consumption (Mauchline, 1980). This is possibly true in Mysis relicta. Borgmann (1978) found that nicotinamide adenine dinucleotide (NADH) oxidation rates were equal to succinate oxidation rates and hence oxidation is also approximately proportional to the rate of oxygen consumption. The rate of production of carbon dioxide by Neomysis integer was measured by Raymont and Krishnaswamy (1968). This was done by determining changes in the pH of the medium. The carbon dioxide output of this species maintained a t 18+0*5"Cranged from 0.10 to 0.62 pl carbon dioxidelmg wet weightlh. General studies of excretion in mysids are few and can be appropriately summarized here. Hargrave and Geen (1968) found that a, mysid (un-named) excreted about, 2.4 pg-at x 10-5 inorganic phosphoruslmg body weightlday. The dominant nitrogenous excretory product is ammonia with lesser quantities of amino-N and apparently no urea (Jawed, 1969). Ammonia amounts to about 75% of the total excreted nitrogen while amino-N represents about 2074. Jawed found that the proportion of nitrogen excreted as ammonia increased at lower temperatures. The average rate of excretion of ammonia a t 10°C by Neomysis rayii wa3 25 pg N/mg body N/day. Chin (1974) found that the rate of ammonia excretion in N . awatchensis changed

8.

151

PHYSIOLOOY AND RESPONSES TO THE ENVIRONMENT

seasonally, average, rates of about 42-58 pg Nfmg body N/day occurring during the summer as compared with rates of 25-42 pg N/mg body N/day during the winter. The amount of total nitrogen excreted at low salinity ( 8 %,) increased with decreasing temperature while the proportion of amino-N increased (Chin, 1976). A similar effect of low salinity and decreasing temperature on the rate of phosphorus excretion occurred. Rates of excretion of ammonia-N were 6-16 pg nitrogen/mg dry weightlday, of amino-N 1-2-58 pg nitrogenfmg dry weightlday, and of total phosphorus 0.2-7.8 pg phosphorus/mg dry weightlday, depending upon conditions of'temperature and salinity. Lee and Chin (1976) found that female Paracanthomysis hispida, over a temperature range of 7-23"C) excreted 2.5-8.0 pg nitrogenlmg dry weightiday as ammonia-N, 1.4-1-9 pg nitrogenfmg dry weightiday as amino-N, whereas comparable rates in males were 4.4-12.6 and 1.9-3.6 respectively. Starvation of the animals significantly decreased the rates of excretion. Raymont et al. (1968) determined the rate of excretion of ammonia-N in terms of body nitrogen in Neomysis integer as approximately 200 pg nitrogenlmg body nitrogenlday. Studies of energy budgets of mysids are few. The partitioning of ingested food between growth, reproduction, respiration, egestion and moulting is shown in Table XXV. The environments of the two species cited are very different. Metamysidopsis elongata is a littoral species living between 35"N and 22"s in the eastern Pacific; the population studied by Clutter and Theilacker (1971) lives in environmental temperatures of 14-20°C. Mysis relicta on the other hand, was studied by Lasenby and Langford in two freshwater lakes, namely Char Lake located in the Arctic at 74" 42") 94" 54'W and the temperate Stony TABLE XXV. PERCENTAGE ENERQY BUDGETS OF MYSIDS Temperature ("C)

Species Metamysidopsis elongata Female Male

14-20 14-20

Mysis relicta, female Stony Lake Char Lake

2-10

0.2-3.8 ~~~~~~~~

G

R

Re

M

36

9 0

49 54

7 10

15.9 11-5

5-7 3.3

63.7 70.2

14.7 14.9

35

~

Ingestion = Growth + Reproduction+ Respiration+ (Moulting+ Egestion), or I = G R + R e +M. Environmental temperature is given. Data from: Clutter and Theilacker, 1971; Lasenby and Langford, 1972.

+

152

THE BIOLOGY OF MYSIDS

Lake at 44" 35'N, 78" 45'W. I n females, the proportional amount of ingested energy ascribed to growth and reproduction decreases with decreasing environmental temperature (Table XXV). This reflects the increased time required for the females to develop at lower temperatures from the egg, attain sexual maturity and produce and lay their own eggs. According to Clutter and Theilacker, the males use a small amount of energy for reproductive products and they assumed it was negligible (Table XXV). The males require less total ingested energy than the females in order t o attain sexual maturity, but expend relatively more of this energy in respirat,ion and moulting. The most important physical parameter of the environment, governing the behaviour and distribution of many species, is the presence or absence of light. Environmental illumination was mentioned previously in the context of various aspects of the animal's physiology. Smith (1970) exposed Hysis relicta to 10 h of normal laboratory levels of illumination during each 24 h period for some 12 weeks; a group of control animals was kept in constant darkness. Survival was high in the control, 93.3%) but only 18.6% of the animals exposed to the periodic illumination survived the experiment. Higher survival rates of many planktonic crustaceans occur under conditions of constant darkness in the laboratory, but it is difficult to relate this result directly to the environment, especially in the context of some of the littoral species examined. Macquart-Moulin (1965, 1972, 1973b, 1977a, b, c) and MacquartMoulin and Patriti (1966) have made the most detailed studies of the behavioural reactions of mysids to light. The phototactic responses of Neomysis americana and Mysis relicta have been examined by Herman (1962) and Beeton (1959) respectively. I n general, mysids are attracted to weak sources of light, but avoid bright light. Bright light often inhibits swimming activity. Normal diurnal light levels of 103-104 lx are inhibitory and produce a negatively phototactic response in most species. This negatively phototactic response was still present at a light intensity of 1 lx in species such as Siriella jaltensis, Anchialina agilis,Gastrosaccus lobatus, G. mediterraneus and G . spinifer. The phototaxis becomes positive a t intensities of 4 x lx or less. Negative phototaxis is present in blue and green monochromatic light of 6 lx, but it is reversed in light of the same intensity but of longer wavelength, as found in Anchialina agilis by Macquart-Moulin (1977b). The 24 h cycle of change in ambient light intensity is the dominant factor controlling the die1 vertical migration of mysids (Beeton, 1960; McNaught and Hasler, 1966; Heubach, 1969; Teraguchi et al., 1975). Mysis relicta migrated towards the surface layers when ambient light

8.

PHYSIOLOOY AND RESPONSES TO THE ENVIRONMENT

153

intensities were decreasing through the range 150-10 lx; the morning descent took place as the light intensities increased through the range 10-3-10-2 lx. Holmquist (1959b) suggests that species such as M . litoralis, which occurs in both freshwater and marine situations, tend to occur deeper in freshwater lakes and a t lower light intensities than comparable populations of the same species in marine habitats. These patterns of responses to changes in environmental illumination result frequently in littoral hyperbenthic species retiring to the shelter of macro-algal beds during the day, where they remain relatively inactive. The nocturnal period of darkness stimulates them to swim and forage outside the beds in the pelagic environment. The earlier work of Foxon (1940) demonstrated this behaviour in various hyperbenthic mysids. The burrowing species of the genus Gastrosaccus usually remain in the sediment during the day but forage in the water column at night (Macquart-Moulin, 1977~).Mysidium columbiae, according to Steven (1961)) appears to be exceptional in that it lives close to the water surface in the tropics, even in the mid-day sun. The positive phototactic response of mysids to weak light sources means that they can be attracted to light lures. Light lures can be used in combination with nets to sample mysids (Brattegard, 1970a; Bacescu, 1975). Brown and Talbot (1972) showed that Gastrosaccus psammodytes were attracted to narrow beams of light in laboratory tanks. This response, however, was absent in the presence of water currents, even weak water currents that did not impede the swimming of the mysids. This die1 pattern of activity affects sampling of mysids, especially hyperbenthic species ; larger numbers often occur in night samples, as instanced by the results of Jansson and Kallander (1968) in sampling Praunus species and Neomysis integer. Polarized light was detected by Mysidium gracile and used for directional orientation (Bainbridge and Waterman, 1957, 1958; Jander and Waterman, 1960). This species tended to orientate its longitudinal axis perpendicular to the plane of polarization. Some terrestrial arthropods have been shown to use linearly polarized light in the sky as a means of navigating during migration but this has not yet been demonstrated in mysids. Environmental temperature affects many of the physiological processes of the animal, as discussed previously. The tolerance of mysids to changes in environmental temperatures varies between species and, to a lesser extent, between populations of the same species in different environments. The freshwater Mysis relicta studied by Smith (1970) in Trout Lake, Minnesota, USA did not tolerate temperature rises, even when these were as small as 1"C/day. Acclimatization

154

THE BIOLOGY OF MYSIDS

of this species to raised or lowered environmental temperatures is probably a slow process and Smith concluded that hypolimnetic temperatures as high as 10°C and epilimnetic temperatures as high as 14°C would adversely affect populations of this species. Holmquist ( 1959b) found that this species in northern Europe and Greenland tolerated a range of O-lS"C, but only if the temperature was kept constant. The vertical and horizontal distribution of temperature in t'he water column was found by Beeton (1960) and Reynolds and De Graeve (1972) to influence the extent of the vertical migration of this species. Thermoclines with gradients of 1.67"C and 2°C acted as barriers to the upward migration of M . relicta; a warm epilimnetic layer was usually avoided although this species is certainly present in such a layer on occasions (Mundie, 1959; Beeton, 1960). Hakala (1978) found that the distribution of M . relicta in Lake Paajarvi, Finland was strictly controlled by the position of the 7°C isotherm, it living a t lower temperatures. The upper lethal temperature for Neom,ysis mercedis (awatschensis)is between 24 and 2505°Cand this species, unlike Mysis relicta, is better able to withstand thermal shocks, even as great as a 14°C rise in temperature (Hair, 1971). Heubach (1969) presents similar results for this species and also points out that Neomysis americana has an upper lethal temperature in this range. The related species, N . intermedia, which occurs through the same range of latitude, is tolerant to a wide range of environmental temperatures, 0-35°C (Murano, 1966d) ; reproduction of this species, however, only takes place between 15°C and 30°C (Murano, 1 9 6 6 ~ ) Neomysis . intermedia may live in freshwater lakes with summer temperatures ranging between about 4°C and 30°C. Very little information on environmental temperatures relating to the distributions of individual species of mysids is available. It is probable, as Kinne (1955) suggests in the case of N . integer, that juveniles have different ranges of tolerance to temperature changes than the adults. This brackish water species was shown by Vlasblom and Elgershuizen (1977) t o be more tolerant of high environmental temperatures than Praunus Jlexuosus ; this is to be expected from comparing the distributions of these two species in coastal regions. The effect of salinity alone, or in combination with temperature, on the physiological processes has been discussed. As with temperature, the tolerances of mysids to changing salinities varies immensely from species to species. McLusky and Heard (1971) point out that Praunus Jlexuosus has been recorded living in varying ranges of salinity, namely 2-18 21.5-27.9 and 35-37-8 They tested the salinity tolerance of this species and found it to range from 2-33 in individuals

x0,

x0

x0.

x0

8.

PHYSIOLOGY AND RESPONSES TO THE ENVIRONMENT

155

from a sea loch in western Scotland. It survived for only 10.75 h at salinities of 0.5 and 50 but tolerated a salinity of 40 for 114 h. Vlasblom and Elgershuizen (1977) found that Neomysis integer was more tolerant of salinities lower than 18 and less tolerant of salinities higher than 18 than Praunus JEexuosus; the latter species tends to inhabit a more bypically marine habitat while the former OCCLUS in brackish and occasionally freshwater habitats. Stepwise did not cause mortality in acclimatization to salinities as low as 21 Leptomysis mediterranea a t 10°C but mortality was present when the salinity was decreased suddenly (Lucu, 1978). Some of the burrowing species of the genus Gastrosaccus may be exceptionally euryhaline. As mentioned earlier, embryo bearing female G. msangii remain buried in the sand, even intertidally after the tide has receded. Consequently, such mysids will be subjected to freshwater seepage, and the effects of rainfall on the beaches. Brown and Talbot (1972) found that G. psammodytes can withstand a 25% dilution of its environmental sea water for at least 42 h with no ill effect. A mortality of 20% took place in 42 h at a dilution of 50%. There was only 30% survival after 42 h in one quarter strength sea water, but survivors recovered completely when returned to normal strength sea water. Fishelson and Loya (1968), on the other hand, found that G . sanctus lives on the Mediterranean shore of Israel in a very high range of salinity of 3638 and could reproduce in even more saline water in the laboratory. Neomysis intermedia, occurring in freshwater and brackish lakes of northern and eastern Japan, tolerates a salinity a0 high as 45 and as low as 0.04-0.06%, (Murano, 1966~).Two species, Mysis litoralis and M . oculata living in the White Sea, were shown by Khlebovich et al. (1970)to be able to tolerate salinities as low as 10 while Holmquist (195913, 1963a) concludes that both these species are euryhaline, M . litoralis certainly occurring in freshwater environments. She further found that the freshwater Mysis relicta could be transferred from fresh water to salinities of 7 and survive there for several months. This predominantly freshwater species occurs in the Baltic Sea in a range of salinity of 3-11.7 and Ackefors (1969) states that it does not occur in regions where the salinity exceeds 12 Very few observations have been made on salinity tolerance of mysids at different environmental temperatures. The tolerance to salinity change is liable to be less at higher temperatures as Vlasblom and Elgershuizen (1977) found in Praunus jiexuosus and Neomysis integer. Likewise, there are few definitive studies of the distributions of mysids in the natural environment in terms of salinity and temperature distributions. A littoral species that appears to be restricted to 8

x0

X0,

x0

x0

x0

x0

x0 x0,

x0.

156

TEE BIOLOGY O F MYSIDS

relatively narrow range of salinities is Metamysidopsis insularis. Quintero and Roa (1973)found that it occurred in littoral mangrove lagoons of Venezuela in a salinity range of 32-7-37.0 Environmental salinity apparently determined its distribution and it did not occur in regions where the salinity was less than 32.7 T u r x r and Heubach (1966)and Heubach (1969)noted that Neomysis mercedis (awatschensis) occurred most commonly in freshwater and brackish regions of salinity 7.5 in the Sacramento-San Joaquin Rivers, USA; population densities decreased in regions where salinities were higher than 18 Neomysis americana however, although occurring in estuaries at salinities in the range 15-20%, (Hulburt, 1957; Cronin et al., 1962) is probably more common outside the estuaries in open coastal situations, including the Georges Bank (Whitely, 1948). Hodge (196313) concluded that Rhopalophthulmus brisbanensis preferred a range of 12.5-33 while Gastrosaccus dakini preferred 5-15 in the Brisbane River, Australia. These salinity preferences appeared to control the seasonal and spatial movements of the populations up and down the river. Ranges of salinity found in the Cananeia region of Brazil did not limit the distributions of Mysidopsis tortonesei and Brasilomysis castroi (Almeida Prado, 1973). Dissolved oxygen concentrations in the water affect the behaviour of mysids to some extent. Childress (1971a, 1975) and Belman and Childress (1976) have shown how Gnathophausia ingens is adapted to live in the oxygen minimum layers of the eastern Pacific. A related species G. zoea, was found to recover even after subjection to anoxic conditions for a period of 4 h (MacDonald and Gilchrist, 1978). NO comparable investigations of littoral or coastal species have been made although some of them are undoubtedly subjected not only to low oxygen concentrations but also to severe changes in concentrations. Ackefors (1969) found that Mysis mixta occurs in regions of the Baltic where the oxygen concentrations are as low as 1-2ml/l water. Holmquist (1959b) found that the freshwater Mysis relicta could probably tolerate levels of dissolved oxygen as low as 29% saturation, or about 2-3 ml O,/l sea water, and Ackefors (1969) suggests that it can live in concentrations down to 1.0m102/1. Other individuals appeared to survive even in the presence of small concentrations of hydrogen sulphide. The behaviour and distributions Of the following species relative to environmental concentrations of dissolved oxygen have been examined : Neomysis mercedis (awatschensis) by Turner and Heubach (1966) and Heubach (1969); Mysis litoralis and M . oculata by Holmquist (1959b); Neomysis intermedia and N . japonica by Murano (1966c, d) ; Metamysidopsis insularis by Quintero

x0. x0.

x0

x0.

x0,

x0

8.

PHYSIOLOGY AND RESPONSES TO THE ENVIRONMENT

157

and Roa ( 1973) ; Bowmaniella brasiliensis, Brasilomysis castroi, Metamysidopsis elongata atlantica, Mysidopsis tortonesei and Promysis atlantica by Almeida Prado (1973). There is no evidence a t present that dissolved oxygen concentrations act as primary controlling factors in the behaviour of the species so far investigated. Variable r a g e s of hydrostatic pressure are experienced by different species of mysids. Bathypelagic mysids are subjected to pressures within the range 100-300 atm and Amblyops magna, caught by Belyaev (1966) at more than 7000m depth, was living at a pressure of some 700 atm. Childress et al. (1978) use a thermally insulated codend on their pelagic trawls to capture Gnathoph-ausia ingens. This cod-end does not protect the animals from the processes of decompression, which they experience during hauling to the surface. This species, however, caught in this manner lives, moults and grows in the laboratory in controlled temperature conditions for many months (Childress and Price, 1978). The effects of the rise in ambient temperature during hauling to the surface are more severe than the effects of decompression, as far as survival of the animals is concerned. Mysids are probably sensitive to small changes of hydrostatic pressure, as Rice (1961) demonstrated in the littoral Schistomysis spiritus. It responded continuously to a cycle of pressure changes comparable to that which it would -experience in its natural habitat on the shore during a tidal cycle. Brown and Talbot (1962), on the other hand, produced no convincing evidence that another littoral species Gastrosaceus psammodytes, responded to changes of the order of 1.5 atm, and Enright (1962) found no response by Archaeomysis maculata to changes to 0.1 atm. Mysids maintained in tanks illuminated from above or below move upwards when the pressure is increased and downwards when it is decreased, the source of the vertical light having little effect. Mysids maintained in horizontally lit tanks respond to increases in pressure by moving towards the bright light or by decreased activity. The species tested by Rice (1967) were Leptomysis lingvura, Praunus jlexuosus, P . neglectus and Schistomysis spiritus. A further species, Siriella armata, behaved differently, the upward movement in increased pressure being more pronounced when lit vertically from above than from below. The tolerance of some littoral species to unrealistic pressures is indicated by the LD,, of Neomysis integer at 160 atm measured by Schlieper (1968). Tidal flows affect the behaviour of littoral and estuarine species. Mysids normally living sub-tidally during daylight are often immigrant in the tidal region a t night (Colman and Segrove, 1955; McLachlan

158

THE BIOLOQY OB MYSIDS

et al., 1979). No endogenous tidal rhythm has been demonstrated in mysids and tidal events appear to modify rather than control aspects of their behaviour. Many littoral species are dispersed between high and low water marks at high tide, but become concentrated a t the water's edge at low tide (Liao, 1951; Matsudaira et al., 1952; Colman and Segrove, 1955 ; Zatkutskiy, 1970; Connell, 1974) ; t,he populations become dispersed again throughout the intertidal area on the following high tide. Flood tides in the Sacramento-San Joaquin River Estuary modify the vertical distribution of Neomysis mercedis (awatschensis); the flood tide has a greater velocity than the ebb tide (Heubach, 1969). This species does not occur in regions of the channel where water velocity exceeds 12cm/s (Turner and Heubach, 1966), a velocity probably equal to or exceeding the maximum cruising speed of the mysid. Mysids orientate in flowing water, facing into the current (rheotaxis). This rheotaxis produces a surprising effect in populations of the burrowing Gastrosacc.tcs psammodytes. Brown and Talbot (1972) demonstrated positive rheotaxis in this species and observed that burrowing takes place as the animals are left dry on the beach by a receding wave. They are thus facing the shore at that time and an examination of the orientation of individuals dug from the foreshore confirms this because they had retained this orientation. Clutter (1969) found that the orientation into water currents by Metarnysidopsis elongata was present under conditions of constant darkness, although less proficient; over 90% of mysids were orientated 15' either side of the direction of the water current when illuminated while only 26% were so orientated in darkness, a further 46% being orientated in the sectors 15' t o 90' either side of the current. Mysis gaspensis was seen by Dadswell (1975) to cling to small pebbles in water currents of 40 cm/s in an estuary in Newfoundland. Observations of the freshwater Mysis relicta from a submarine, by Robertson et al. (1968), showed that it appeared to cling to a hard substrate where possible and were orientated into the current; these observations were made at a depth of approximately 42 m. Wave action in the littoral region affects the distribution of many intertidal species (Boroditch and Havlena, 1973 ; Brown and Talbot, 1972). It tends to depress their vertical range on the shores, the mysids moving offshore to deeper water to avoid the effects of breaking waves. Breaking waves also wash Gastrosaccus species out of their intertidal burrows. Consequently, mysids respond to changes in the great majority of the physical and chemical factors of their environments but the detailed responses can vary between species. This reflects the wide

8. PHYSIOLOGY

AXD RESPONSES TO TEE E ~ O N M E N T

159

variety of environments colonized by mysids. These responses to changing environments govern the distribution of many coastal species present within localized areas. They often make location of the populations and interpretation of the results of sampling a sea area difficult. Mysids respond to one another and their social behaviour also influences their distribution within a sea area. The social behaviour of mysids is discussed in the following chapter.

CHAPTER 9

BEHAVIOUR The changing physical and chemical factors of the environment, as discussed in the last chapter, can therefore enable mysids to occur in alternative parts or areas of the environment at different times. This is most noticeable in coastal and especially the littoral and intertidal environment. Vast numbers of a species can be sampled in a restricted area one day, and be completely absent from that area on the following day. Consequently, quantitative sampling of the populations has to take into account the whole local horizontal and vertical range of the species. This is most easily done by recognizing a coastal area that has a number of relatively discrete environments such that each species of mysid is almost completely restricted to one habitat. A relatively isolated, sheltered bay or inlet in an otherwise rocky coastline usually provides suitably isolated and contained populations of intertidal species. River estuaries or fjords are other areas where populations of a species can usually be easily located and monitored. It is important in studying populations of a species in such an area t o know the locations of other populations of this species in adjacent areas. An estimate can then be made, in the context of the physical, chemical and topographical characteristics of the region as a whole, of the possible amount of communication (immigration and emigration) between the populations under study and those in the adjacent areas of the region. This is necessary to distinguish real from apparent increases in population numbers. Real increases result from active breeding and production of young, while apparent increases can result from one or both of the following : an immigrant population from an adjacent area moving into the area and mixing with the resident population or a disaggregated population aggregating or swarming in a small sea area, so that it is sampled more effectively. The converse of these two processes, emigration and disaggregation, can take place and the result is an apparent decrease in the population numbers. The sizes of relatively isolated populations of coastal species were estimated seasonally in various regions of the west coast of Scotland and 58’N by Mauchline (1971f). The between latitudes 55’30”

J

M

M

J

S

N

J

M

M

J

S

N

700 400 300

200

8000

I 120

8 brasi/iensis

I

4000

40

1600

;$oo

800

60

20

FIG.44. Seasonal occurrence of species of mysids. Anchidim agilis: histogram, Lochs Etive and Creran (Mauchline, 1971f); dotted line and right scale x 2, Gulf of Marseille (Macquart-Moulin, 1965); hatched line and right scale, Plymouth (Tattersall, 1938'. BowmanieEZa brasiliensis, Cananeia Region, Brazil (Almeida Prado, 1973). Brasilomysis castroi, Cananeia Region, Brazil (Almeida Prado, 1973). Erythrops elegans : histogram, Clyde Sea area and Loch Ewe (Mauchline, 1971f); dotted line and right scale, Gulf of Marseille (Macquart-Moulin, 1965). E. serrata, Clyde Sea area (Mauchline, 1971f) ; Qastrosaccus dakini, Brisbane River, Australia (Hodge, 196313). Q. normani: histogram, Clyde Sea area and Loch Ewe (Mauchline, 1971f); hatched line and right scale, Plymouth (Tattersall. 1938). G. spinijer: histogram, Loch Ewe and Gruinard Bay (Mauchline, 1971f); dotted line and right scale, southern North Sea (Boysen, 1977); hatched line and right scale x 10, Kiel Bay, Baltic Sea (Baan and Holthuis, 1971). Hemimysis Zamornae Clyde Sea area (Mauchline, 1971f). Black histograms, ovigerous females and mature males ; cross-hatching, other stages.

J

M

M

J

S

N

FIG.45. Seasonal occurrence of species of mysids. Leptonzysis grucilis : histogram, Clyde Sea area (Mauchline, 1971f); dotted line and right scale, Gulf of Marseille (MacquartMoulin, 1965); hatched line and right scale x 3, Plymouth (Tattersall, 1938). L. lingvura: histogram, Clyde Sea area and Loch Ewe (Mauchline, 1971f); hatched line and right scale, Isle of Man (Liao, 1951). Mesopodopsis slabberi: histogram, southern North Sea (Baan and Holthuis, 1971); dotted line and scale x 10, Gulf of Marseille (Macquart.-Moulin, 1965); hatched line and scale x 0-1, Kiel Bay, Baltic See (Boysen, 1977). Metumysidopsis elongata, California (Fager and Clutter, 1968). M . elongata ntlnntica, Cananeia Region, Brazil (Almeida Prado, 1973). Mysidium columbiae, Jamaica (Goodbody, 1965). Mysidopsis angusta, Clyde Sea area (Mauchline, 1971f). M . clidelphys, Clyde Sen area (Mauchline, 1971f). M. gibbosu, Clyde See area Mauchline, 1971f). M. tortonesei. Cananeia Region, Brazil (Almeida Prado, 1973). Black histograms, ovigerous females and mature males; cross-hatching, other stages.

J

M

M

J

S

N

800

400

4000 2000

3000

I N mercedis

1000

40

20

10000

tc

f?

1SO?

6000

flexuosus

i

u

800 2000 FIQ. 46. Seasonal occurrence of species of mysids. My& mixta, Kiel Bay, Baltic Sea (Boysen, 1977). Neomysis americunu, Indian River Inlet, Delaware (Hopkins, 1965). N . mercedis, Sacramento-San Joaquin River Estuary, California (Heubach, 1969). N . integer: histogram, Loch Etive (Mauchline, 1971f); hatched line, Baltic Sea (Kinne, 1955). Paramysis arenoaa, Clyde Sea &ma and Loch Ewe (Mauchline, 197Lf). P . baceacoi, Roscoff, France (Labat, 1957). P . nouveli, Roscoff, France (Labat, 1957). Pseudomma aflne, Clyde Sea area and Loch Etive (Mauchlinj, 1971f). Praunua Jexuosw,: histogram, Clyde Sea area and Loch Etive (Mauchline, 1971f); hatched line, Denmark (Blegvad, 1922). Black histograms, ovigerous females and mature males ; cross-hatching, other stages.

J

M 1

1

M 1

1

J '

S 1

1

N 1

J 1

~

M

M

J

S

N

1 1 1 " 1 1 1 1 1 1

~

s spirirs 2000

1200

400

120

100 20

40 750

250

R brisbanensis

30

100 10

400

FIG.47. Seasonal occurrence of species of mysids. Pruunus inernzis: histogram, Clyde Sea area (Mauchline, 1971f); hatched line, Denmark (Blegvad, 1922). P . neglectus: histogram, Clyde Sea area (Mauchline, 1971f); hatched line and right scale, Denmark (Blegvad, 1922). RhopuZopthaZnaus brisbunensis, Brisbane River, Australia (Hodge, 1963b). Schistomysis kervillei : histogram, Loch Ewe (Mauchline, 1971f); hatched line, southern North Sea (Baan and Holthuis, 1971). S. ornuta: histogram, Clyde Sea area and Loch Etive (Mauchline, 1971f); hatched line and right scale, Plymouth (Tattersall, 1938). S. spiritus: histogram, Clyde Sea, area and Loch Ewe (Mauchline, 1971f); hatched line, southern North Sea (Baan and Holthuis, 1971). Siriella cZTmUtU: histogram, Loch Ewe (Mauchline, 1971f); hatched line, Gulf of Marseille (MacquartMoulin, 1965).S. cluuaii: histogram, several areas of west coast of Scotland (Mauchline, 1971f); hatched line and right scale, Gulf of Marseille (Macquart-Moulin, 1965). S. @ensis: histogram, several areas of west coast of Scotland (Mauchline, 1971f); dotted line and left scale, Gulf of Marseille (Macquart-Moulin, 1965); hatched line and right scale, Isle of Man (Liao, 1951). S. norwegicu, Clyde Sea area and Loch Etive (Mauchline, 1971f). Black histograms, ovigerous females and mature males; crosshatching, other stages.

9.

BEHAVIOUR

165

occurrences of these species are shown in Figs 44-47 along with the occurrences of other species in other latitudes investigated by Labat (1957), Hodge (1963b), Goodbody (1965), Hopkins, (1965), Fager and Clutter (1968), Heubach (1969), Baan and Holthuis (1971), Almeida Prado (1973) and Boysen (1976). I n addition, Moore et al. (1979) record seasonal changes in the numbers of Neomysis integer present in the Severn Estuary while Hakala (1978) studied the changing number of Mysis relicta in Lake Paajarvi in southern Finland. All species, with the possible exception of the tropical species Mysidium columbiae (Fig. 45), show seasonal periods of maximal and minimal occurrence. The populations of M . columbiae studied occurred at 19"N in Jamaica. Southern sub-tropical species occurring in latitudes 25-27'5, namely Bowmaniella brasiliensis, Brasilomysis castroi, Gastrosaccus dakini, Metamysidopsis elongata,atlantica, Mysidopsis tortonesei and Rhopalophthalmus brisbanensis, all show seasonal maxima and minima of population numbers. Mauchline (1971f) divided the species of the west coast of Scotland into groups according to their seasons of maximal occurrences. These groups have been re-examined in the context of whether the individual species are near their northern or southern limits of geographical distribution, or occur in coastal, littoral or estuarine habitats. The seasonal maxima of occurrence of the Scottish species (Mauchline, 1971f) are listed in Table XXVI, along with the babhymetric habitat in which they were studied; the latitudinal range of the study area is 55"30'N to 58"N. Other species with the latitudes in which they were studied are also listed. No obvious patterns are present in the data and only one aspect is worth mentioning in some detail. The maxima of occurrence of littoral and estuarine species are in the spring, summer and autumn with minima in the winter except in Erythrops elegans (Fig. 44) which occurred maximally in the winter and spring. Gastrosaccus spinifer, a littoral species in the west of Scotland, was found to occur maximally during the winter (Fig. 44) by Boysen (1976) in the southern North Sea and by Baan and Holthuis (1971) in Kiel Bay in the Baltic Sea; the populations studied by these authors were not littoral but neritic. Leptomysis lingvura in a littoral habitat in the Isle of Man (Liao, 1951) has a winter maximum of occurrence (Fig. 45). Mesopodopsis slabberi in a littoral habitat (Macquart-Moulin, 1965) has summer and autumn maxima but in neritic habitats (Baan and Holthuis, 1971; Boysen, 1976), has autumn and winter maxima of occurrence (Fig. 45). Schistomysis kervillei in an offshore habitat (Baan and Holthuis, 1971) has a winter maximum as does 8.spiritus (Fig. 47). Estuarine and littoral species, therefore, tend to occur maximally during the warmer months of the year, but if they occur in offshore

TABLEXXVI. SEASONAL OCCURRENCE AND AGGREGATION OF MYSIDS Most interne breeding periods species

Study area

Species range . Habitat 58N-25N

C

Bowmankllu b r a s i l h s k 25s B r a a i h y s b CaetrOi 25s Erythrops ebgans 58N

25N-20s 23s-25s 58N-25N

L L L

No (Yes 42N) Yes Yes Yes?

E . serrclta Gaatrosmua abkini a. n m n i a. spinijeer

56N 27s 58N 58N

?ON-50N 275-35s 58N-8N 59N-8N

C E C L

Yes Yes ?(Yes 42N) ?(Yes 54N)

Hemimysis l a n a o w

56N

57N-41N

C

Yes?

LeptornysiS grccCiliS L . liTlgVura Mesopodopsb a j k m M . skbberi

56N 56N 32s 54N

58N-25N 58N-25N

-

C L E L

Yes Yes Probably No (Yes 42N)

Metmn.yysidop& elowata M . elongata atlanticr;C My+urn wlumbiae MysidopsG anguata M . didelphys

32N 25s 19N 56N 56N

35N-22s 255 24N-llN 68N-31N 68N-40N

37N-335

of

occurrence

B r d throughout ( W , winter ;Sp, spring; the year Su, summer;A , autumn)

56N

Anchialina agilie

Seasonal mima

L L

ges Yes Yes

C

No

of

Type oj aggregation

W

Breeding aggregation inW A Shoals su Shoals .W, Sp Shoals, breeding aggregation in Sp Sp, Su, A Shoals Sp, Su Shoals W? unknown Sp, Su Breeding aggregation in sp , Sp, SulA Breeding aggregation in Sp and Su W Shoals Sp, Su, A Shoals Su, A Swarms Su, A Shoals and swarms

L L

Source

SP P Sp, Su, A? Sp, Su, A W W ? ?

Swarms and schools Swarms Shoals and schools Unknown Breeding aggregation

inw

10 11 11 10

10 4 10 10 10

10 10 12 1, 13 8 11 3, 6 10 10

M . gibboaa M . tortonesei My& rnixta N m y & amm'cana N.W V d i S N . integer Paramy& arenoaa P . baceswi

P . rwuveli Praunua $exuoaua

P. immk P.neglectus Pae-mu

aflm

s.o m t a s.82yiritu-9 Siriella armata s.clauaii S. j d t e d 8. norvegica

56N 255 38N 38N 56N 58N-56N

58N-29N 23s-25s 78N-40N 5ON-35N 59N-35N 66N40N 57N-32N

46N 46N 56N 56N 56N 56N

50N-42N 50N-28N 58N-40N 58N-48N 68N-48N "ON-40N

21s

54N

L L C C

Yes Yes

Sp, Su, A

A

?

I

Yes Yes Yes Yes

Sp, Su Sp, Su Sp, SU,A Sp, SU

Sp, Su, A Shoals Sp,A Swarms

10 11

unknown

Su, A Su, A Su, A Sp, Su, A

13 5, 7

C

Yes Yes No Yes No No

275

E

Yes

Shoals Shoals Shoals and s w a m Shoals, breeding aggregation in Su Su, A su Shoals Su, A Su, A Shoals Sp, Su, (A) Sp, Su, A Shoals Sp, SU Sp,Su,A Shoals Sp,Su(A) Sp, Su,A Shoals W W Breeding aggregation inW Sp, Su, A SulA Shoals

58N

58N-49N

L

Yes

Sp, Su

Sp, Su

Shoals and swarms in late Su

10

56N

59N-25N

C.

Yes

W

W, Sp

Breeding aggregation

10

Yes Yes ?(Yes42N)

Sp, Su(A) Sp, SU ?

inW Sp, Su, A Shoals and s w a m Sp, Su,A Unknown W unknown

10 10

3 ?

? ?

W W

10 10

58N 58N 58N 58N 56N

59N-28N 58N-30N 58N-32N 58N-27N 58N-27N

L E L L L

L L

L

L L C C C

1

Unknown unknown

9 10

10 2 2 10

10 10

10 4

10

"he latitude of the study areas 21given along with the latitudinal range of the species. "he habitat 2I shown ~8 estuarine (E), Littoral (L) and neritic (c).Breeding Strategy noted for the study area and for 81'088 south of it in parentheses. Sources of data: 1. Blegvad, 1922; 2, Labat, 1957; 3, Steven, 1961; 4, Hodge, 1963b;5, Rioharde and Riley, 1963; 6, Goodbody, 1965; 7,Hopkins, 1965;8,Clutter, 1969;9. Heubach, 1969;10,Mauchline, 1971f; 11, Almeida Prado, 1973;12,Connell, 1974; 13,Boysen, 1977; 14, Wooldridge, 1977.

168

THE BIOLOGY OF MYSIDS

pelagic or neritic environments the seasonal maxima of occurrence tend to occur in the autumn and winter. The neritic species are listed in Table XXVI. Nine of the thirteen species have winter maxima. Three of the remaining four species, Hemimysis lamornae, Mysis mixta and Neomysis americana, range from the littoral to the neritic environment. The fourth species, Erythrops serrata, is strictly neritic. The reason for such a difference between the majority of species or populations of species occurring in the estuarine and littoral environments on the one hand and those occurring in the neritic or coastal environment on the other, is not known. Baan and Holthuis (1971), collecting mysids from the surface plankton near the “Texel” lightship in the southern North Sea, noted this seasonal pattern of occurrence of winter maxima, but could not explain it. They discuss the possibilities of seasonal and reproductive migrations. Such a migration has only been demonstrated within the restricted habitat of a Scottish sea loch; age groups of #chistomysis ornata are distributed within the loch as shown in Fig. 21, p. 78 and an ontogenetic horizontal migration is performed seasonally. One possible explanation of this pattern is that several neritic species have been found to form breeding aggregations. Such aggregation of a previously disaggregated population will result in large samples from a study area that is also a breeding area of the species. Most neritic species breed in the late winter and spring and so aggregation of the populations for purposes of breeding would be present at these seasons. The process of aggregation or disaggregation of a population of mysids can be detected in two types of sampling programmes. The first programme entails a large series of samples taken throughout a region. The period of sampling should extend over one year to describe the cycle within the populations, but sampling can be restricted to shorter periods to describe the existent states of the populations. Samples should be small in that each sample is taken from relatively small volumes of water or the sampler should be towed over 25 or 50 m serial lengths of transects in littoral regions. Mauchline (1971f) used a D-net incorporating a 75 cm diameter tow-net, a conventional 75 cm diameter tow-net, and a 1 m beamtrawl (for neritic species). Other suitable samplers are shown in Fig. 18, p. 67. The environments sampled extended through and beyond the local distributional ranges of each of the species studied. Individuals of a disaggregated species can be expected to occur in samples in small numbers, even as single individuals, while individuals of aggregated species can be expected to occur in larger numbers or not at all in samples. The aggregations and swarms of a species in a

9.

BEHAVIOUR

169

localized area will not contain all individuals of that species occurring in that area. Some inditriduals do not conform to the general behavioural patterns of the majority of individuals in the population. Steven (1961) suggests that about 5 % of the population of Mysidium columbiae remain independent of the shoals of this species. This proportion of non-conformist animals probably varies at different times within and between populations of species. The overall result, however, is that sampling in areas where dense shoals and swarms of a species occur usually produces three types of samples: the first has large numbers of animals, the second none and the third has single or few individuals. The result of such sampling programmes on the west of Scotland are shown in Table XXVII. Taking Siriella species as examples, S. armata and S. clausii occur more commonly in samples in numbers ranging from 2 to 50 while S. jaltensis occurs more commonly as single individuals. Consequently, it is probable that the former two species tend to be sampled from aggregations while S. jaltensis, although sometimes caught from aggregations, is more commonly sampled from disaggregated populations. The numbers of other species in samples, for example Leptomysis gracilis, Neomysis integer, Praunus jiexuosus and Schistomysis spiritus, show bimodal distributions, the species tending to occur in samples in either small or very large numbers. A similar analysis of samples of mysids collected on the Caribbean coast of Colombia was made by Brattegard (1973); his dat’a have been supplemented with data from his other papers and re-analysed (Table XXVIII) to conform with the information on the Scottish species (Table XXVII). A comparable analysis of the samples of mysids, many of them oceanic species, described by Tattersall (1955) from the “Discovery” collections is given in Table XXIX. The interpretation of the data on a few of these species is more difficult. The samplers used range from 50 cm through 4.5 m tow-nets to large rectangular nets. Consequently, the volume of water filtered when taking individual samples probably ranges over several orders of magnitude. The types of nets in which each species was predominantly caught are indicated in Table XXIX. The four species Eucopia australis, E . grimaldii, E. sculpticauda and Gnathophausia gigas were caught by large nets, The rest of the species were taken predominantly in nets of comparable size to those used to catch species listed in Tables XXVII and XXVIII. The da%ain Tables XXVII-XXIX suggest %hatthe great majority of species aggregate, a t least during certain periods of their life cycles. The seasonal maxima and minima of population numbers in Figs 44-47 reflect real increases and decreases in the sizes of the populations as well

170

THE BIOLOGY OF MYSIDS

as aggregation and disaggregation within the populations. Increases in the sizes of samples taken .of populations of most of the littoral species investigated are a result of simultaneous addition of juveniles, through reproduction, and aggregation of both the juveniles and the adults. Aggregation within different populations of mysids can be very different in origin, form and function, and the following terms were defined by Mauchline (1971f):

Aggregation. A population is aggregated when individuals are not randomly distributed throughout the area or volume in which they are found ; this area or volume may be a neritic bathymetric zone parallel to the shore. No inference is made in using this term to the factors, for example physical or chemical parameters of the environment or behavioural reactions between individuals, that might be responsible for this uneven distribution. Breeding aggregation. A breeding aggregation is a grouping together, in a non-random distribution, of sexually mature individuals for the purpose of mating. Restricted distribution. A population has a restricted distribution if it occurs in a defined part or several defined parts of a sea area or volume. The inference here is that environmental parameters are probably responsible for the uneven distribution of the populations and that behavioural reactions between individuals are less important in maintaining the aggregation. Shoal. Clutter (1969) defines shoals as large groups of mysids, ranging from a few metres to tens of metres across, that occur within their habitat zones (areas). He uses this term in the context of species that live in shallow water, probably in open bays. Shoals however, can also occw in more strictly pelagic habitats. Swarm. According t o Clutter, swarms are smaller, more integrated groups of mysids than those comprising shoals. Swarms are often constituent parts of shoals. The term swarm implies greater cohesiveness but not parallel orientation of individuals. School. Like swarms, schools can be constituents of shoals (Clutter, 1969), but schools differ from swarms in consisting of polarized groupings of individuals swimming in the same direction. Wittmann (1977) does not use the above terms as defined by Mauchline (1971f) but re-defines some of them primarily on a geometrical and secondarily on a social basis. The terms are used throughout this discussion in the form defined above. A detailed knowledge of the biology, life history and local distributional patterns of a species is required before the types of aggregation

TABLEXXVII. THENUMBERS OF SAMPLES WHICH CONTAINED THE STATED NUMBERS OF INDMDUALS OF EACH SPECIESOM THE WESTCOASTOF SCOTLAND (after Mauchline, 1971f)

Total nos of Species

1

Anchialina agilis 5 Erythrops elegana 23 E . serrata 14 Bastrosaccus n o r m n i 5 0. spinifer 3 Hemimysis lamornue 13 Leptomysis yracilis 14 L . lingvura 19 Mysidopsis anguata 19 M . didelphys 33 M . gibbosa 20 Neomysis integer 7 Paramysia arenosa 8 Praunus fiexu~aus 18 P . inermis 19 P . neglectua 18 Pseudommu a&e 7 Schistomysis kervillei 9 S . ornuta 46 S. apiritus 11 Siriella armatu 1 S . clauaii 2 S . jaltensis 7 S . norvegiea 3

2-5

6-20

21-50

51-100

101-200

6 15 32 1 5 14 23 15 25 35 31 8 18 29 27 21 6 6 39 11 6 3 3 1

2 4 29 1 4 0 20 9 4 23 34 4 26 23 23 25 5 2 33 8 5

1 4 21

1 1 7

1 10

2 2

2 2 6 2 6 11 10 12 19 19 12 1 4 15 2 1 2

1 2 1

3 1

201-500

501-1500

>1500 Samples

5

23

25

1

3 9 5 9 8 11

2 2 8 8 5 9

2 29 2 19 8

5 12

3

7

1

5 4 3

3 5 7

2 1 13

1

7

6

15 48 118 7 14 38 117 47 48 102 114 86 83 121 124 76 19 32 143 68 13 7 12 6

Indiu-idu.uk 148 407 4222 13 170 276 32890 420 117 1107 7540 32133 3122 7399 9435 775 114 2307 2388 31081 107 64 33 28

Mean Number number of indiof viduals indiin VidWllsJ largest sample sample 10 8 36 2 12 7 281 9 2 11

66 373 38 61 76 10 6 72 17 457 8 9 3 5

65 108 305 6 49 72 1664 122 12 151 930 2871 247 44 1 1623 48 33 599 248 3635 34 25 9 14

TABLEXXVIII. THE NUMBERS OF SAMPLES WHICH CONTAINED THE STATEDNUMBERS OF INDMDUALS OF EACH SPECIES FROM THE CARIBBEANREGION

Total nos of Species Amuthimysis cherados A. gibba A. polita Anchialina typica Antromysis bahamensis Boutanidla d i s s k i l i s B. parageia B. sewelli Brmilom?ysiseastroi Cubanmysis jimenezi Dwptromysis paucispinosa D. spinosa Erythrops parva

1

2-5

6-20

6 9 7 13 21 13 2 8 6 11 14 10 2 4 1 3 2 5 4 3 1 2 8 9 5 5 1 3 7 1 5 7 7 1 2 3

2 5

3 4

21-50 2 5 7 4 5 1

51-100

101-200

201-500

501-1500 ~ 1 5 0 0Samples

2 3

4

3 5

3 3 8 1

4 4

5 1

1 2

1 2

2 1

2 2

2 1

-

1 2

2

1

1

1

-

Individwzls

24 54 26 45 39 12 6 25 41 20 16

175 532 520 2723 8607 91 33 163 4073 968 69

11 18

494 1029

Nean Number number of indiof widwzls indiin viduals/ largest sample sample 7 10 20

6 7 99 48 4

39 82 57 1376 3200 44 19 40 1624 227 30

45 57

184 684

61

221 8

Heteromysis elegans

1

2

H . mayana

5 2 4 5 5 4 2 1

5

Metamysidopsis swifti Mysidella minutu Mysidiurn colurnbiae M . gracile M . integrum Mysidopais almyra M . ankeli M . bispinulata 1 M . brattstroemi 4 M. cultrata 3 M . furcu 2 M . mathewsoni 8 M . mortenseni 3 M . robusta 4 M , taironana 2 &. tortonesei 7 M . velifera 9 M . virgulata Promysis atlantica 3 Siriella chierchiae '73

2 4 8 2 2 3 1 1 3 1 1

7 3 6 3 2 15 3 2 18

5 3 8 8 2 10 1 2 1 5

2 -

1

3 6 1 2

5 3 4 11 4 4 18

5 1 2

10

-

1

8

6

2

5

1

1

1 2 4

2

1

-

2

1

-

-

1

1 4 2 1 4

6 2 6 2 5

1 -

10

1

8

2 1 5

1 6

0

I

2

3

>

Y

~~

Data from: Brattegard, 1969, 1970a, c , 1973, 1974a.

9 19 48 11 27 6 5 7 16 5 6 20 10 10 11 28 49 17 13 77

120 23 455 220 6455 113 2211 20 68 178 201 14 326 90 71 20 344 14190 1020 1135 433 4188

12

2 51 12 134 10 82 3 14 25 13 3 54 5 7 2 31 507 21 67 33 54

37 5

357 48 1472 51 804 11

35 59 48 7 170 18 28 5 262 3899 125 174 122 1321

TABLEXXIX. THE NUMBERSOF SAMPLES WHICH CONTAINEDTHE STATED NUMBERSOF INDIVIDUALS OF EACH SPECIES OF OCEANIC MYSIDS

Total nos of species

Netsa

1

2-5

6-20

21-50

1 18 4 6 9

2 34 3 4 3 13 10 17 7 8 11 6 7 12 16

2 32 1

3 12 4

4 1 7 1 1

1

51-100 101-200

201-500

+ + +

Y Y Y Y Y+ Y Y Y Y+ LY LY LY

19 8 27 9 7 16 20 21 11 12

2 4 1 8

2

2

1 5 3c

4

Individuals

~

~

Anchialina truncata Antarcknnysk maxim A . oh.linii Arachnmysis megalops Boreomyais brucei B. rostrata Caesaromysk hkpidu Dactykmblyop8 hodgsoni Euchaetmera intemedia E . tenuis E . typica E . zurstrmseni Eucopia australis E . grimldii 1. seulpticauda

>500 Samples

Mean Number number of indiof viduals indiin viduals/ largest sample sample

15*

1

9 120 14 11 12 38 19 53 17 16 27 28 34 24 36

204

23

70

26 18 358 44 201 34 38 41 59 133 59 141

2 2 9 2 4 2 2 2 2 4 2 4

11 3 240 6 26

6 6 4 11 44 13 20

E . unguiculata Gnathophausia gigas G. ingens Hansenomysis falklandica Katerythrops oceunae Longithorax capensis Lophogaster challengeri Meterythrops picta Mysidetas d i m r p h a M . macrops M . microps M . patagonica M . posthon Mysidopsis acuta Pseudamma armatum P . calmani P . sarsii SiriZllu thompsonii

Y L Y + Y Y + Y +

++

+ + +

1

5

6

8 22 5 9

9 10 4 3 2 3 7 2 3 3 2

7 3 5 3

-2

6 7 4

+

-

+ + Y

4 6 17

1 5

2 6

-

12

1 1

1

3 1

27 17

114 33

4 2

34 18 1' 11 7

63 192

2 11 1 4 3 3

1'

1 5 0 1

3 4 4

5

3 1 3 2 7

9

1

2

2 4 5 1

1

1 3 2

1

'.

1

1 1

1

9

1

9 13 9 10 27 26 9 10 10 37

17 41

24 30 154 414 540 76 >400 379 270 36 16 175

17 3.' 60 8 15 15 30 4 4 5

21

5 6 47

3 23 11 5 70 92 300 39 >lo0 9s 107 13 3 36

Data from: Tattersall, 1955. a ,Samples in small nets, less than 1 m diameter; Y, young fish trawl (TYF and TYF 70B); L, 2-4.5 m tow-nets, large rectangular nets. b Quantity described as hundreds. Described as many.

+

176

THE BIOLOGY OF MYSIDS

formed can be defined. Consequently, little can be said about the nature of the aggregations of the'oceanic species listed in Table XXIX. The samples of Tattersall (1955) and others, suggest that juveniles, as well as the adults, of many of these species aggregate and it is probable that social behavioural mechanisms in concert with tile physical-chemical parameters of the environment act to maintain these aggregations. The aggregations of the Scottish species (Table XXVII) were studied in the context of the biology of each of the species (Mauchline, 1971f). Some species such as those of the genus Siriella, Gastrosaccus normani and Mysidopsis angusta were rare and the factors involved in their aggregation remain unknown. Breeding aggregations, consisting predominantly of sexually mature adults occurred at certain times of the year in populations of Anchialina agilis, Erythrops elegans, Gastrosaccus spinifer, Hemimysis lamornae; Mysidopsis didelphys, Paramysis arenosa, Pseudomma ajfine and Schistomysis ornata (Table XXVI). The four species, Erythrops elegans, Gastrosaccus spinifer, Hemimysis lamornae and Paramysis arenosa are littoral and usually permanently located near or just below low water mark, even a t high tide. The other species are strictly neritic living to depths as great as 200 m. The populations of these eight species never attain the size and densities reached by populations of strictly littoral species, the mean numbers of individuals per sample ranging from 6 to 38 (Table XXVII). Two of these species, Erythrops elegans and Paramysis arenosa, in addition form shoals. Shoals are also formed by Erythrops serrata, Leptomysis gracilis, L. lingvura, Mysidopsis gibbosa, Praunus jiexuosus, P. inermis and P. neglectus. The mean numbers of individuals of these species per sample ranged from 9 to 281 (Table XXVII). Two of these species, Erythrops serrata and Leptomysis gracilis, are neritic at depths of 30-150m. Very large samples, greater than 500 individuals, of L. gracilis have been caught and it is possible that this species forms localized swarms rather than shoals. Of the Scottish species, only Neomysis integer, Schistomysis kervillei and S . spiritus are known t o form swarms and these can be observed visually in shallow coastal regions. The mean sample size of these species ranged from 72 to 457 (Table XXVII). The most detailed studies of aggregation of mysids have been made on littoral and intertidal species for the obvious reason that they are usually accessible for visual observation. Social behaviour in conjunction with the physical-chemical discontinuities of the environment appear to act together in the formation and maintenance of shoals of several species. Dadswell (1975) found that the entire population of Mysis gaspensis in a Newfoundland estuary formed a shoal 0.1-0.5 m

9.

BEHAVIOUR

177

wide and some 200 m,in length along the eastern side of the estuary at each succeeding low tide. The mysids were dispersed at other stages of the tidal cycles so that they occurred more or less evenly over the entire estuary at each high tide. Dadswell suggests that the shoal is formed a t low water because the mysids select a localized region of the estuary whexe medium water velocities in combination with low salinities occur and these are the factors initiating and maintaining the shoal. This aggregation is then in the category of a restricted distribution rather than a shoal. Whiteley (1948) studied the “large swarms’’of Neomysis americana occurring on the Georges Bank. The greater portion of the Bank has depths in the range 40-100m although it shallows to 5 m depth in its north-western regions. The “swarms” are not swarms as defined above but conform to the definition of shoals. These shoals on the Bank are elongated in shape and occur on the long axis of the Bank within the 75 m depth contour. The species does not occur in numbers outside the 100 m depth contour and this situation indicates a restricted distribution over, howevex, a relatively large area. Separate shoals occur within this area, approximately 20 000 km2, as indicated by the variation in numbers per sample. The samples, however, were taken over a distance of some 1850 m so that little can be deduced about the sizes of individual shoals, nor is there any information on the influences of the physical-chemical parameters of the environment upon the distribution of the shoals. Whitely did find, however, that the average numbers of this species, when it occurred in his samples, were 43.3 animals/m3 water during the day compared with 0-2 animals/m3of water during the night. This situation suggests that dispersion of the shoals takes place at night, probably in the vertical plane through vertical migration. It is common for intertidal species to be “collected” by the receding tide, so that the individuals become concentrated near low water mark (Liao, 1951; Matsudaira et aE., 1952; Colman and Segrove, 1955; Zatkutskyi, 1970; Connell, 1974). Clutter (1967, 1969) observed another aspect of the intertidal, littoral and immediately sub-littoral environment. This is the tendency for many species of mysids to occur in well-defined bathymetric zones roughly parallel to the shore. This coastal zoning extends seawards to depths of 20-30 m in Californian and Scottish waters. Species living in such zones have a relatively, and often rigidly, restricted distribution such that the populations occur in a series of shoals. This is the case with Neomysis integer, Praunus species and Schistomysis spiritus, for example, in Scotland, Metamysidopsis elongata, according to Clutter ( 1967), in California, Gastrosuccus sanctus in Israel (Moran, 1972) and Leptomysis species in the northern

178

THE BIOLOGY OF MYSIDS

Adriabic Sea (Wittmann, 1977). Shoals of Metamysidopsis elongata moved within their zone of occurrence, but there was no correlation between these movements and changing temperature, salinity, light intensity or composition of the substrate. Clutter, however, considered that their movements might be in response to changing concentrations of detrital organic matter. Certainly Mysidopsis gibbosa in the Clyde Sea area is frequently found associated with the area of fallout of terrigenous organic matter on the sea bottom from freshwater streams. Hobson and Chess (1976) found that Siriella pacijica was numerous in shallow depressions in the sea bottom where flocculent organic material had collected. Swarms of Leptomysis species in the Adriatic Sea remained in constant topographical situations, returning to them after disturbance. Wittmann ( 1977) concludes that swarms of different species that occur in overlapping bathymetric zones usually associate with different substrates, while species preferring the same substrate usually occur in different batihymetric zones. Some species are associated with beds of seaweeds or with coral reefs and this association of a species with a substrate, leading to a restriction of its distribution, is discussed further near the end of this chapter. Light has been shown to influence swarm formation (Clutter, 1969; Macquart-Moulin, 1973a; Zelickman, 1974). Swarms of Neomgsis mirabilis persisted from dawn to dusk but dispersed at night fall while those of Metamysidopsis elongata did disperse to some degree during darkness. Dispersion of hyperbenthic populations takes place at night in species that perform a die1 vertical migration. Wittmann (1977) found that at sunset swarms of Leptomysis species expanded, the distance between individuals increasing. Aggregations of different species mixed in the hyperbenthic layer during the night but separated once again at dawn. Wittmann marked individuals in swarms and found that, although the swarm returns to the same topographical region of the bottom at daylight, the individual need not necessarily do so but may have joined an adjacent swarm. Consequently, there appears to be considerable exchange of individuals between swarms of these Leptomysis species. Clutter (1 969) describes the shoals of Metamysidopsis elongata as consisting of schools and swarms. Juveniles appeared to occur in separate swarms from older individuals as Emery (1968) found in populations of Mysidium gracile and Wittmann (1977) found in populations of Leptomysis lingvura. Green (1970), however, describes mixed swarms of Acanthomysis sculpta, and some littoral species form mixed swarms in western Scotland. The shapes of swarms can vary. Clutter found small spherical swarms of about 1000 individuals of an

9.

179

BEHAVIOUR

Acanthomysis species while schools of this species travelled through the water as elongated bodies. Large swarms are often wafer-shaped and about 5 cm in thickness. The distance between individuals in a swarm varies between species and also between size or age groups within a species. The mean distance to the nearest six neighbours in swarms or schools of Acanthomysis species A, was 1.1-2-7 cm (equivalent to about 3 4 times body length), of Metamysidopsis elongata 3.7 cm and of Mysidium columbiae about 4 cm (Table XXX). Clutter estimated the TABLEXXX. NUMBERS AND DENSITIES OF MYSIDS IN SWARMS AND THE MEAN DISTANCE FROM ONE INDIVIDUAL TO ITS NEAREST SIX NEIGHBOURS (Steven, 1961; Fishelson and Loya, 1968; Clutter, 1969)

Species

Numberlswarm

Number/cms

Acanthomysis sp. A

13 16 22 127 137 e3200

0.1 0.3-0.7 0.3 0.1 0.05 -0.1 % 1'

Qarrtrosaccua sanctus Metamysidopsis elongata Juveniles Mysidium columbiae a

-

14000

-

0.02

M e a n distance (cm) 2-1 1.1-1.5 1.5 2.1 2.7 -2.1 0.22 2-4 3.7 4

In sediment at a density of 18/cm2.

mean distance between individuals as the cube root of the mean volume of water occupied by one individual in a pelagic population. Fishelson and Loya (1968),however, estimated the density of Gastrosaccus sanctus in the sand of a beach on an areal basis. Assuming a more or less even distribution of individuals within the swarm, the mean distance between them is the square root of the area occupied by one animal. There are probably threshold densities within an aggregation that trigger swarming and schooling. This is extremely difficult to study in the laboratory situation, but Clutter ( 1969) estimated such densities to be in the range 0.01-0.70 animals/cms. The minimum density required before Metamysidopsis elongata orientates into schools is probably of the order of 1 mysid/l6 cm3, equivalent to L distance between individuals of about 2.5 cm. Schools can consist of as few as 16 M . elongata, or conversely, of many thousands. Clutter found that

180

THE BIOLOGY OF MYSIDS

small swarms or schools did not disintegrate even when disturbed and could avoid towed nets in a co-ordinated and organized manner. According to Zelickman (1974, rain storms cause swarms of Neomysis mirabilis to disperse, but they reform within a period of 60-90 min after the rain storm has ended. Clutter (1969) and Zelickman (1974) experimentally examined the factors such as population density, illumination and water flow that might affect formation and maintenance of swarms. The main conclusion is that vision is important in the organization of swarms, as Steven (1961) concluded fiom his studies of Mysidium columbiae. Chemical cues and mechanoreception, however, are probably also important, especially in maintaining the relatively high degree of association of individuals that is present in shoals and swarms throughout the hours of darkness. This must be accentuated in species such as Gastrosaccus sanctus which, according to Moran (1972)) probably forms shoals at night, individuals of this species being buried in the sand during the day. Pseudomma ufline has vestigial eyes yet populations of this species form breeding aggregations (Table XXVI). Fage (1925) describes the cavernicolous Spelaeomysis servatus, whose eyes are also extremely reduced, as swimming in large shoals near the surface. Hansenomysis falklandica is yet another species with reduced eyes that probably aggregates (Table XXIX). Consequently, aggregation is probably achieved by several or all of the following processes rather than by any one in isolation. (a) The innate drive towards aggregation, triggered in some species by attainment of a physiological condition, e.g. approaching sexual maturation. (b) The physical-chemical discontinuities and gradientis of the environment in conjunction with the physiology of the species resulting in the population having a restricted distribution : this situation produces populations in which the threshold densities that trigger the formation of swarms and schools occur. (c) Visual cues in the presence of threshhold densities. (d) Chemical cues in the presence of threshhold densities bhat may be an order or several orders of magnitude less than those densities active for visual cues. (e) Mechanoreception by the individuals, probably assuming greatest importance in the processes of maintaining and structuring schools and swarms once formed. The relative importance of any one of the above processes in producing shoals, swarms and schools will depend to a considerable extent upon the species concerned and the habitiat in which it lives.

9.

BEHAVIOUR

181

Wittman (1977) found tihat some species such as Anchialina agilis, Siriella jaltensis and S. armata are not active swimmers and associate strongly with a substrate. These species did not form swarms although his descriptions of their occurrence conform with aggregations within restricted distributions. Other species, such as Leptomysis apiops, are active swimmers but aggregate a t biogenic structures ; they also thus have restricted distributions. A third group of species, examples of which are L. lin,gvura, and L. mediterranea, are active swimmers but still show preferences for certain substrates so restricting their distributions ; these species, however, are active swarmers. A final group, instanced by Mesopodopsis slabberi, show no preference for a substrate, are active swimmers and form schools and swarins. The functional aspects of aggregations of mysids are discussed by Clutter (1969) and Zelickman (1974). One probable function is that shoaling aids in maintaining the population more effectively within a zone of occurrence. This applies particularly t o littoral and sublittoral species that may be subject t o longshore or offshore drift. This would also apply t o estuarine species, especially those that occur in channels with marked tidal flows. Neomysis integer shoals close t o mud banks, in an area studied by Kinne (1955), and probably experiences lower water current velocities as Dadswell (1975) found in the case of a shoal of Mysis gaspensis in a Newfoundland estuary. The social interactions of shoaling would reinforce the environmental parameters in maintaining the integrity of the aggregation. The simultaneous release of all young from the marsupium by a female within a shoal results in the young being potential members of the shoal. The swimming speeds, however, of juveniles and adults of a species differ considerably with the result that a mobile shoal becomes a self-sorting mechanism separating age groups and so producing swarms of individuals of closely similar body size (Clutter, 1969). Some species are more active swimmers than others and the occurrence of mixed age groups within shoals of a species may be indicative of a more lethargic life style. The function of breeding aggregations are self-evident in a three-dimensional fluid environment. This type of aggregation appears t o assume greater importance in the lives of the neritic, and probably offshore pelagic, species than in littoral, intertidal and estuarine species. The only littoral species thought to form a conventional breeding aggregation is Acanthomysis sculpta studied by Green (1970). Shoaling is a more or less permanent feature of the populations of shallow water species and so mating is easily achieved; the primary function of shoaling in these species is probably not for breeding. Shoaling in neritic species, on the other hand, does not seem, in many cases, t o be a permanent state

182

THE BIOLOGY OF MYSIDS

within the populations, although many of them have restricted distributions that residt in densities of animals approaching those that occur within shoals. Densities in populations with restricted distributions are probably within the range 0.01-1.0 individuals11 compared with densities of 50-700/1 in swarms (Table XXX). The mean distances between individuals in these restricted distributions would be within the range 10-50cm compared with distances of 1-4cm in swarms. Wittmann (1977) observed Anchialina agilis, a species that does not form swarms in the Adriatic Sea, clinging to Zostera leaves; the individuals were spaced a t distances of about 3 cm. Clutter (1969) suggests that males detect females receptive to mating at distances of about 5 cm. Consequently, breeding aggregations of a species developing within populations with restricted distributions would produce aggregation with distances of this order between individuals. Shoaling undoubtedly affords protection to individuals and populations against predators (Emery, 1968;Clutter, 1969).Swarming appears to be primarily a behavioural trait of shallow water or surface living crustaceans, these being more or less continuously subjected to visual detection by potential predators. Contacts with an isolated mysid or with a swarm of mysids are probably equivalent events and scored as a single detection by the predator. I n considering a population of mysids as a whole, this results in the frequency of detection being an inverse function of the number of mysids in the swarm. Attacks on swarms by fish often cause the swarms to fragment into smaller swarms that immediately move away from the point of attack, only a few individuals being successfully captured by the fish. The tendency for dispersal of swarms a t night, possibly associated with nocturnal feeding of the mysids, suggests that the avoidance of diurnal predation may be an important, if not primary, benefit conferred on the population by this behaviour. The possible significance of swarming in mechanisms of population control and regulation is briefly discussed by Clutter (1969) but much further information is required before any useful conclusions can be drawn. Mysids exhibit the conventional escape reaction of shrimp-like crustaceans ; that is, they flex the abdomen ventrally and anteriorly and are propelled backwards away from the source of danger. Brown and Talbot (1972) found that Gastrosaccus psammodytes produced this reaction not only when swimming pelagically, but also when buried in its burrow in the sand. A disturbance of the water-saturated sand in the vicinity of its intertidal burrow when the tide has receded caused the mysid to leap “tail first into the air”. It landed a few centimetres from

9. BEHAVIOUR

183

its burrow and immediately buried itself once again in the sand. Gastropod snails elicit this reaction from the mysids when they crawl over the sand close to a burrow. Many mysids can change colour quite remarkably by the expansion or contraction of large chromatophores distributed over the body and appendages. The chromatophores are in three main groups. The neural group is associated with the cerebral ganglia and nerve cord, the visceral group with the gut, hepatopancreas and gonads and the caudal group with the telson and uropods. Tattersall and Tattersall (1951), quoting the detailed investigations of Keeble and Gamble (1903, 1904), have described the system in some detail. The chromatophores contain a large amount of dark brown (red in some genera) pigment and a yellow or whitie (sometimes blue or blue-green) reflecting substance. Keeble and Gamble (1904) describe the histology, development and reactions of the chromatophores to light and changing environmental background colours in detail. The early work of Koller and Meyer (1930) and Kropp and Perkins (1933) demonstrated that eye stalk extracts of Mysis stenolepis, Praunus JEexuosus and P. inermis contain hormones that activate the chromabophores. Praunus JEexuosus is translucent when captured over sand, but dark in colour when captured among seaweed. The related species P . neglectus is dark green or brown, dependent upon the colour of the seaweeds in its habitat. Mysidopsis gibbosa is dark in colour when it lives sub-littorally in the zone of fallout of dark terrigenous material from a freshwater stream. Bacescu (1975) briefly discusses the colour variation occurring in littoral Tanzanian mysids. The range of cryptic coloration varies between species but is limited in many. Zelickman (1974) states that the colour of Neomysis mirabilis changes rapidly as a function of locomotory behaviour. Undisturbed animals in the sea are usually transparent! with a yellowish-grey colour. Their colour darkens a few seconds after capture and, under subsequent quiescent conditions, their normal colour is resumed some 30 min later. Mauchline and Webster (unpublished) examined the time required for several species to change colour when placed against a new background colour. Newly caught Neomysis integer, Praunus jlexuosus, P. inermis and P. neglectus require up to 1 h to adopt a new coloration, a situation noted by Chicewicz (1952) in P. jlexuosus. The time required, however, for the mysids to effect a second change of colour, immediately following the first, is much longer, many individuals not achieving it even after 12 h. This suggests that species adopting cryptic coloration may be relatively static. Cryptic coloration would then have two effects: protection against predators and relative restriction of the animals to a

184

THE BIOLOGY OF MYSIDS

given substrate. A species such as P. neglectus, that lives on several adjacent but different Substrates, would tend to remain spread among the substrates and so inhabit a larger area and volume of the environment. There is evidence that some hyperbenthic species of mysids prefer certain types of substrates, as instanced above by the preference of Mysidopsis gibbosa for areas of fallout or settlement of terrigenous material. Boysen (1976) examined the substrates with which Gastrosaccus spinifer, Mesopodopsis slabberi and Mysis mixta are associated in the Baltic and concludes that only the last species shows a preference, occurring on a mixture of sand and mud. The substrate preferences of a range of western Atlantic species are described by Wigley and Burns (197 1). Bowmaniella portoricensis, Mysidopsis bigelowi and Promysis atlantica prefer sand. Neornysis americana typically associates with sand that is free of silt, while Erythrops erythrophthalma lives over sand that contains small to moderate quantities of silt, but not enough to classify the sediment as silty sand. Two species, Boreomysis tridens and Pseudomma afine, associate with silty sand. Amblyops abbreviata and Bathymysis renoculata live on fine sediments, consisting of fine sand, silt and clay. Only one species was associated with a coarse substrate ; Meterythrops robusta occurs on gravels and other coarse materials. Some of these substrate preferences will result in restricted distributions. The spectrum of particle size of sand may be a limiting factor in the colonization of beaches by burrowing species of the genus Bastrosaccus. Brown and Talbot (1972) found that juvenile G. psammodytes burrowed more frequently into sand with a particle size range of 0.55-0-70 mm in diameter than into sands consisting of particles of larger or smaller sizes. Brattegard in his papers on Caribbean mysids, notes the substrates on which the majority of species were captured; many species appear to show preferences. Many sub-littoral species associate with algae, even with specific kinds of algae, and with corals. Mukai (1971) found mysids that were not identified sheltering among the thalli of Sargassum serratifolium in the Inland Sea of Japan. Xiriella longipes and an Acanthomysis species are commonly associated with the Zostera marina belt in Kyushu, Japan (Kikuchi, 1967). Siriella pacifica and Acanthomysis sculpta live within the California kelp beds (Hobson and Chess, 1976). Praunus neglectus is commonly associated with Fucus and Laminaria beds on the coasts of western Scotland. Coral reefs also provide sheltered areas colonized by mysids. Bacescu (1975) states that Mysidopsis coralicola occurs in areas of sand among corals and algae along with Anisomysis maris rubri, A . sirielloides, Gastrosaccus msangii, Siriella brevicaudata

9.

BEHAVIOUR

185

and Tenagomysis tanzaniana., Mysidium species occur among the reefs of the Florida coast of the United States (Emery, 1968) and Alldredge and King (1977) found species that they did not name to be important constituents of the fauna of the Great Barrier Reef, Australia. Coral reefs and algal beds shelter mysids in every case where they have been searched for ; the individual species are probably distributed between separate microhabitats and substrates rather than occurring in mixed populations. The burrowing behaviour of Gastrosaccus species has been studied in some detail. Macquart-Moulin (1977a, c) has examined the die1 rhythms of activity of several species, that is their habit of remaining quiescent in the burrows during daylight hours but emerging a t night to live pelagically and forage for food. Gastrosaccus psammodytes was found by Brown and Talbot and G. sanctus by Fishelson and Loya (1968)to be able to burrow into sand submerged in water and also into water saturated sand in contact with the air. Burrowing is effected by the thoracic legs, both the endopodites and exopodites, sweeping the sand away from the ventral regions of the thorax to form a depression. The abdomen and telson are moved from side to side with a downward pressure so that they become buried in the sand. There is some forward motion, but of limited extent, during burrowing and all movement ceases once the animal is completely covered with sand. The antennae and antennules seem to have no function in burrowing. Burrowing appears to be initiated immediately the thoracic legs come in contact with sediment, Brown and Talbot describing the reaction as a thigmokinesis. Emergence from the burrows normally takes place as the following high tide floods the beach with water. Some individuals, through the action of the thoracic legs, emerge from the sand spontaneously while others are washed out by wave action. Moran (1972) found that G. sanctus is not evenly distributed in burrows over a beach in Israel; the population is concentrated in a zone of occurrence parallel to the water’s edge. Gastrosaccus species are probably not the only mysids that bury themselves wholly or partially in the sediments. The neritic Lophogaster species may beha.ve in this way. Robertson et al. (1968) observed the freshwater Mysis relicta from a submarine and found that some individuals partially buried themselves in mud at depths of 20-100 m. Some species of mysids live in commensalassociation with other organisms. This commensalism is assumed rather than clearly demonstrated, except in the case of Heterornysis actiniae and the sea anemone Bartholomea annulata. Randall et al. ( 1964) observed Mysidium gracile move to shelter in the spaces between the spines of the echinoderm

186

THE BIOLOGY OF MYSIDS

Diadema antillarum Philippi when danger threatened. Emery (1968) found that schools of Mysidiurn gracile, when threatened by a predatory fish, crowded into the nesting caves of several species of pomacentrid fish that inhabit reefs. The nesting fish made no attempt to eat the mysids, but drove off the potential predator. Other conimensal associations are shown in Table XXXI. The best investigated association is TABLEXXXI. COMMENSAL ASSOCIATIONS OF MYSIDS Mysid

Associated organism

Antromysis anophelinae Land crab Cardisoma guanhumi Anemone Bartholomea annulata Heteromysis actiniae (Leseur) H . gerlachei Gastropod shells of hermit crabs H . gomezi Cavity of a sponge Ophiuroid Astroboa nigra Doderlein H . gymnura H . harpax Gastropod shells of dardanid crabs H . mariarci Cavity of a sponge Gastropod shells of dardanid crabs H . odoiatops H . zeylan ica Cavity of sponge Siphonochalina siphonella Levi Idiomysis tsurriarnali Anemone Megalactis hemprichi Medusa Cassiopea andromeda Spelaeomysis Land crab Cardisoma guan.humi card isomae

Reference

Bowman (1973) Clarke (1955) Tattersall (1967) Bacescu (1970b) Tattersall (1962) Tattersall (1962) Bacescu (1970b) Clarke (1955) Tattersall (1967) Bacescu (1973a) Bacescu (1973a) Bowman (1973)

that between Heteromysis actiniae and the sea anemone Bartholomea nnnulata. Clarke (1955) states that this mysid swims in small groups among the expanded tentacles of the anemone but does not appear to be attacked by the nematocysts. It rests for periods on the basal portions of the tentacles of the anemone where there are fewer arrays of nematocysts. Some B. annulata have no cream coloured crescentic array of nematocysts present on these basal regions of the tentacles and, according to Clarke, the mysids recognize and prefer these individuals. The mysids feed on the undigested food ejected by the anemone. Another mysid, Heteromysis bermudensis, may also occur in association with the anemone (Clarke, 1955). Further investigations of other species, especially in the genus Heteromysis, may provide further examples of commensalism.

CHAPTER 10

POPULATION DYNAMICS The numbers of individuals in populations of most species of mysids fluctuate seasonally in a regular manner. A contributory cause of such fluctuations can be changing degrees of aggregation or disaggregation within the populations as discussed in the last chapter. Such changes in the spatial distributions of individuals can result in apparent increases and decreases in population numbers. The major cause, however, of these seasonal fluctuations is the seasonal pattern of reproduction of the population. Determination of these patberns is often difficult, especially in the cases of neritic and oceanic species. Most investigations so far made have been confined to populations of species living in restricted environments that can be sampled comprehensively. A more or less standard method has evolved for studying the population dynamics of a population of mysids. It involves sampling the population successively ab intervals of one, two or four weeks, the sampling interval depending upon the rates of change in the cornposition of the population being studied. The individuals witihin bhe population can be ascribed to one of the following categories: (a) juveniles, secondary sexual characteristics are absent ; (b) immature males, the secondary sexual characteristics are in process of development ; (c) mature males, the secondary sexual characteristics are fully developed and the gonads are normally functional ; (d) immabure females, the marsupium is developing and is smaller in size than that of the mature female, no young are carried ; (e) mature female, with the marsupium fully developed, but not yet filled with young ; (f) ovigerous females, mature females with young present in the marsupium ; (g) empty females, females with a marsupium from which the young have emerged, these females have not moulted after releasing the young.

188

TEE BIOLOGY OF MYSIDS

All of these categories, are useful in investigating many species. There are species, however, where one of these categories, especially the last, is either missing or of little value in the investigation. The sex of a juvenile cannot normally be determined by an examination of its external morphology. The secondary sexual characteristics that develop are different in different genera. All females have a marsupium that develops in stages a t successive moults and the pleopods may be normally developed or vestigial. The males may or may not have penes, one or more pairs of pleopods fully developed or specially modified, or an enlarged antennular peduncle armed with setae. The form of the secondary sexual characteristics of any one species can be determined from the appropriate descriptive literature listed in Mauchline and Murano (1977). The presence of these characteristics in the process of development is used to ascribe the individual to the category of either immature male or female. The mature males are recognized by the full development of the secondary sexual characteristics and its state of sexual maturity can be confirmed quite easily by examining the gonads. Consequently, the first four categories listed above are relatively easy to recognize. The fifth category, mature females with a marsupium present, but not filled with embryos is more difficult. This category is unimportant in most species studied, because the full development of the marsupium is associated with egg laying. It occurs, however, in a species such as Praunzls jlexuosua in Scotland. The female can release an autumn brood of young and immediately moult to a form with a large marsupium that is not filled with embryos. She is a mature adult that has bred a t lea& once but does not breed again until bhe spring of the following year. This category was not used by Mauchline (1971~) in a study of the biology of this species because it is difficult to distinguish from immature females. They can sometimes be distinguished by their longer body length from immature females with a welldeveloped marsupium. There are species, usually offshore oceanic or neritic species, that may release their embryos prematurely on capture. This results in a larger proportion of “empty females” in the sample than might be expected. This is the last category listed above which, in practice, is usually recognizable. The lamellae of the marsupium are usually distended, thin-walled and irregularly disposed. It$appearance is markedly different from that of the immature or mature female discussed above, that have compact, regularly arranged lamellae. The division of the individuals in a sample among several or all of these categories allows an assessment of the state of the population t o be made. It is important, however, to ensure that the youngest juveniles are being effectively sampled. The size of these juveniles can be

10. POPULATION

189

DYNAMICS

determined by an examination of the oldest larvae, those with stalked eyes, that occur within the marsupia. The youngest juveniles swimming freely in the sea will have a body length that does not exceed that of these larvae by more than 25%. The next step is to measure the sizes of the individuals in the different categories. Body size is usually determined as a measurement of total length: this measurement can be made from either the base of the eyestalk or the apex of the rostrum t o the posterior end of the telson and uropods (excluding the setae); a further measurement, standard length, (Grossnickle and Beeton, 1979) is used and is the distance between the apex of the rostrum and the posterior edge of the sixth abdominal segment. Carapace length has been used by Ishikawa and Oshima (1961), Matsudaira et al. (1952), Murano (1964a), Amaratunga and Corey (1975) and Pezzack and Corey (1979) as an index of total length. The length of the uropod was used by Clutter and Theilacker (197l), total length in Metamysidopsis elongata being equal to 4.5 times the length of the uropod. Total length is the accepted measure of body size in mysids and most, but, not all, authors who have used other indices have defined the relationship of the index used to total length. Such relationships are given in Table XXXII along with TBLE XXXII. REGRESSION CONSTANTSOF BIOMETRICAL RELATIONSHIPS Regression constants y = bx+a or log y = bloga: Species

Parameters

&

a ~ t ~ o s a cvulgaris cus

TL on CL

- 0.538

Myeidopsis didelphys

CL on TL

0.434

juveniles

0-203

females

0.647

- 0.048

males Myaia stenolepis juveniles

Reference

Matsudaira et al. (1952) 0.242 Mauchline (unpub1ished) 0,264 Mauchline (unpublished) 0-230 Mauchline (unpublished) 0-269 Mauchline (unpublished) 3.935

TL on CL 0.037

adult females Praunw inermis

b

+a

2.191

CL on TL

0.051

Amaratunga and Corey (1976) 7-963 Amaratunga and Corey (1975) 0.212 Mauchline (unpublished)

3.034

contd.

TABLEXXXII-contd. Regression constants y = bx+a or log y = b log x Species

M etamysidopsis elongata

Parameters log, WD on log, TL

BD on TL Boreomysis arctica maturing females

a

- 5.436 0.031

b 2.77 0.117 3.42

spent females

- 3.74

3.98

adult males

- 1.68

2.31

log WD on log TL

- 4'5654 2.00

Stony Lake

- 4.4622 2.84

Lake Michigan

-4.1549

2.92

- 1.27 - 1.58 0.66

2-84 3.42 1-10

Finland, < 3 mm CL log WAF,, on log CL on log CL >3mmCL log W, Neomysis americana TL on SL April-September

(log WD + 1)on log SL

- 0.353

0.741

October-March

(log W,

+ 1)on log SL

- 0.598

0.975

log W, on log TL

-4.569

2.691

log WD on log TL

-3.914

2.723

Neomysis m.irabilis

0.018

2.3

Paramysis intermedia

0.002

3.4

Paramysis kowalewski

0.002

3.0

Mesopodopsis slabberi

0.012

2.5

Acanthomysis strauchi

0.020

2.5

Paramysis baeri

Reference Clutter and Theilacker (1972) Clutter and Theilacker (1972)

log WD on log TL

- 2.92

Mysis relicta Char Lake

+a

log W, on log TL

Brattelid and Matthews (1978) Brattelid and Matthews (1978) Brattelid and Matthews (1978) Lasenby and Langford (1972) Lasenby and Langford (1972) Reynolds and De Graeve (1972) Hakala (1978) Hakala (1978) Richards and Riley (1967) Richards and Riley (1967) Richards and Riley (1967) Shushkins et al. (1972) Shushkina et al. (1972) Nekrasova and Rakitina (1968) Nekrasova and Rakitina (1968) Nekrasova and Rakitine (1968) Nekrasova and Rakitina (1968) Nekrasova and Rakitina (1968)

BD, body diameter; CL, carapace length; SL, length measured from tip of rostrum to base of telson ; TL, total length; W,, body dry weight; WW,body wet weight; WAFDW, ash-free dry weight.

10.

191

POPULATION DYNAMICS

those between wet and dry weight and total length. The relationship between antenna1 scale length on the one hand and total and standard lengths on the other are shown graphically for Mysis relicta by Grossnickle and Beeton (1979)but no regression equations are given. A graph relating body dry weight to carapace length in Neomysis intermedia is given by Murano (1964a).Boroditch and Havlena (1973)have published two graphs relating body wet weight to t o h l length in Paramysis intermedia and P . ullskyi. Clarke (1962) measured various parts of the body in four Gnathophausia species (Table XXXIII) ; the lengths of all parts are regressed TABLEXXXIII. REGRESSIONCONSTANTS O F BIOMETRICAL RELATIONSHIPS OF THE BODY AND TOTAL BODYLENGTHIN GNATHOBETWEENVARIOUSPARTS PHAUSIA SPECIES ~~~~~~~~~~~

~

~

Regression constants y = bx+a Species

G . ingena

$+$?

Independent variable CL

cw AL TeL

UL

0. gigas

G . zoea

6+$?

6 ?

6 ? 6+?

RL ASL 70L CL

cw

AL TeL UL RL ASL CL CL AL

AL TeL

UL RL ASL 70L

n 233 229 234 234 234 147 233 28 55

54 58 58 58 37 58 26 22 26 22 48 48 47 48

20

a

-2.4378 0.3276 2.3201 0.5877 -0.2194 12.4393 0.2073 -42.3813 -1-6915 0.3050 0-5158 0.0780 -0.1977 11.4368 1.1871 -1.5758 -0.6897 0.1764 0-4802 0.7512 0.8152 11.3070 3.0234 -22.7793

b

0.4795 0.1677 0-3530 0.2225 0.1750 0.0736 0.1243 0.4628 0-4283 0.1435 0.4063 0.2375 0.1948 0.1345 0.1432 0.4693 0.4995 0,3822 0.3744 0.2026 0.1783 -0.0141 0-0932 0.4629

Regression coescient r

0.9980** * 0*9975*** 0.9981* * * 0*9977*** 0.997 7** * 0.7579*** 0.9951 * * * 0*9551*** 0.9987 * * * 0.9958*** 0*9986*** 0.9973*** 0*9959*** 0-7357*** 0.9945*** 0.9975** * 0*9977*** 0.9910*** 0.9963*** 0.9947*** 0*9928*** - 0.2579 0.9638*** 0.9802* ** contd.

192

THE BIOLOGY OF MYSIDY

TABLEX X X I I I - c o n t d . Regression constants y = bz+a Species

G . gracilis

3

0 d+P 6 ? d+P

G. gracilis (dwarf)

d+? d ? S+P

Independent variable CL CL

cw AL AL TeL UL EL ASL 70L CL CL

cw

AL AL TeL UL RL ASL 70L

Regression coeficient

n

a

b

r

77 130 183 76 131 208 207 110 206 89 19 33 45 19 33 51 52 36 52 33

- 1.1413 -2.3284 -0.2013 0.5441 1.9813 10.2364 -0.2317 6.2667 0.9194 - 25.8196 -0.0461 -3.3350 - 1.6371 1.5013 2.8512 0.3876 9.9140 11.8026 1.1555 - 18.5246

0.3912 0.4130 0.1530 0.4310 0.4019 0.0705 0.2053 0.1623 0.1558 0,4123 0.3704 0.4464 0.1843 0.4127 0.3716 0-1975 - 0'0039 0.0487 0.1450 0.4940

0.9969*** 0*9976*** 0.9965*** 0*9980*** 0.9976*** 0.5362*** 0.9967*** 0-8606*** 0*9949*** 0.9176*** 0*9833*** 0.9896*** 0.9839*** 0*9941*** 0.9879*** 0-9816*** - 0.1597 0*2294*** 0-9608*** 0.9634***

Data from: Clarke, 1962. Independent variables : AL, abdomen length, ASL, antennal scale length; CL, carapace length; CW, carapace width; RL, length of rostrum; Tel. telson length; UL, uropod length; 70L, length of 7th brood lamella (oostegite).

on body length, measured from the base of the eyestalk to the posterior end of the telson excluding the setae. Carapace length is measured from the base of the eyestalk to the posterior lateral edge excluding any spinous process; abdomen length is measured dorsally from the anterior edge of the fist t o the posterior edge of the sixth segment; telson and uropod lengths are measured from the posterior edge of the sixth abdominal segment to the posterior edge of the appendage excluding the setae; antennal scale length is that of the scale alone; the length of the rostrum is from the base of the eyestalk to the anterior tip of the rostrum. No sexual differences were present in the measurements except in those of the carapace and abdomen relative to total length in G . zoea and G. gracilis; separate equations have been calculated for these relationships (Table XXXIII). The scatter within the raw

10.

193

POPULATION DYNAMICS

data is very limited as can be seen in Fig. 48 and deduced from the high values of the correlation coefficients. All the correlation coefficients are significant a t 0.1 yo level except those indicated. The rostrum is subject to damage.

8

20-//:. -30

..:'ES .:
,,..il'-'

10-

0

./"

TG;.z-:e

.,,*rp-'

a

. ::!*H,:~c_.,::;..*ddo-U

4

a v

d,&72*--

-10 a

-3:"-

I

I

1

g c

-20

i---

ropod length

1

-..+ ...." :::... ,,.:

< 2

.

'*o;:' :.

I

I

I

I

I

I

1

I

I

Fro. 48. Regressions of the dimensions of various parts of the body on total body lengt h in Qnathophausia ingens. The respective regression equations are given in Table XXXIII. Data from: Clarke, 1963.

The size of the lith of the statocyst is used by Morgan and Beeton (1978) in analysing the occurrence of individual instars within a

population of Mysis relicta. The frequency distribution of the diameters of the liths was polymodal and the modes appear to correspond with successive instars, although interpretation of them involves the usual difficulties of polymodal analyses. They found that the diameter of the lith ranged from 0.04 to 0-26 mm; their measurements were accurate to 0.01 mm. A decrease in the size interval with a consequent increase in the number of size classes might aid in interpreting the modes, especially among size classes of the smaller individuals. Studies of the populations of a range of species from different latitudes have been made (Table XXXIV). The proportions of different

TABLEXXSIV. POPULATION STUDIESOF MYSIDS IN DIFFERENT GEOGRAPHICAL REGIONS 0bservations Species

Latitude

Anchialina agilis

Region

56'30" 43ON Boreomysis nobilis 70"N Bowmuniella brasiliensis 25"N Brasilomysis castroi 25's Erythrops elegans 56% Erythrops serrata 56"N Gastrosaccus brevifissura 32% Gastrosaccus dakini 27'30's 43'N G . lobatusla. normani Gastrosaccuspsammodytes 35"s Gastrosaccus sanctus 46"N 38"N Gastrosaccus vulgaris Gnathophausia ingens 38"N-32"N

Scotland Marseille W. Greenland Brazil Brazil Scotland Scotland S. Africa Australia France S. Africa Black, Azov Seas Japan E. Pacific

Leptomysis gracilis Leptomysis lingvura

Scotland Scotland, Irish Sea

Mesopodopsis africana Mesopodopsis slabberi

56"N 58"N-5 6"N

32"s 55"N 43"N 46"N Metamysidopsis elongata 33"N M . elongata atlantica 25"s Mysidium columbiae 18"N Mysidopsis bigelowi 38"N Mysidopsis didelphys 56'N Mysidopsis gibbosa 56"N Mysidopsk tortonesei 25"s

S. Africa Denmark France Black, Azov Seas E. Pacific Brazil Jamaica W. Atlantic Scotland Scotland Brazil

Breeding periods

few seasonal

X

A W(?) S P SU A a Su Sa sp su a w sp su a w S P SU a w(?) SPSUaw SPSUAa SPSUAa SPSUAw sp su a w SP SU A sp su a

X

?

2(?) 3 0.12

X

S P SU A w SPSUAw

2 2

X X

X X X X

X X X

X X X

x X X X

X X X X X X X X

Generations1 Year

sp su a a SU A SP SU A a SP SU A SP S U A w sp su a w sp su a w sp SU a AW S P SU A w sp su a w

2 ?

2-3 2 ?

?

2 2 >3

2 1 3

References Mauchline (1971e) Macquart-Moulin (1965) Dunbar (1940) Almeida-Prado (1973) Almeida-Prado (1973) Mauchline (1968) Mauchline (1968) Connell (1974) Hodge (1963b) Macquart-Moulin (1965) Brown and Talbot (1972) Zatkutskiy (1970) Matsudaira et aZ. (1952) Childress and Price (1978) Mauchline (1968a) Liao (1951); Mauchline (1969a) Connell (1974) Blegvad (1922) Macquart-Moulin (1965) Zatkutskiy (1970) Fager and Clutter (1968) Almeida-Prado (1973) Goodbody (1965) Hopkins (1965) Mauchline (1970b) Mauchline (1970b) Almeida-Prado (1973)

75"N-72'N 42"N-4OoN 74'N 74"N

Arctic W. Atlantic Laptev Sea Char Lake

63"N-61°N 61"N 49ON-47'N 46'-43'N 45ON

Great Slave Lake L. PiiiijiLrvi, Finland L. Superior L. Huron Ontario Stony Lake

42'N Mysis stenolepis Neomysis americana

My& Mysis Mysis Mysis

litoralis mixta oculata relicta

SP w ( ? )

-

0.5? 0.5 0.5?

X

SP

0.5

X

X

SP W SP SU or M7 SP

L. Michigan

X

sp su a w

1

45ON-42"N

W. Atlantic

X

SP w

1

38% 41°N

Delaware, USA Long Island Sound

X X

sp SU a sp SU a w

Neomysis mercedis

45'N 42"N-40°N 38"N

Passamaquoddy Bay Georges Bank E. Pacific

Neomysis integer

56ON-54"N

Nwmysis intermedia Paramysis arenosa Paramysis bacescoi

X X

X

X X X

0.5 0.5- 1.0 0.5 0.7? 1

2-3 2(?)

-

X

sp su a S P SU A (w?) sp su a w

Scotland, Baltic

X

SPSUAw

2-3

54"N 54'N

West Ireland Netherlands, Baltic

X

X

S P SU A usp su a

2-3 3

40°N 58ON-54"N

Japan Scotland, Irish Sea

X

SP su A w SPSUAw

2-3 2-3

X

S P SU A w

2

Roscoff

X X

X

3

3 2(?)

Geiger (1969) Wigley and Burns (1971) Geiger (1969) Lasenby and Langford (1972) Larkin (1948) Hakala (1978) Carpenter et al. (1974) Carpenter et al. (1974) Lasenby and Langford (1972) Reynolds and De Graeve (1972); Morgan and Beeton (1978) Wigley and Burns (1971); Amaratunga and Corey (1975) Hopkins (1965) Richards and Riley (1967) Pezzack and Corey (1979) Wigley and Burns (1971) Turner and Heubach (1966); Heubach (1969) Kinne (1955); Mauchline (1971b) Parker and West (1979) Vorstman (1951); Wiktor (1961) Murano (1964a) Liao (1951); Mauchline (1971a) Labat (1957)

cont.

TABLEXXXIV-contd. Observations speciw

LatitwEe

Paramysis intermedia

Region

few seasonal

Russia

Breeding PETiods

Generations/ Year

SP SU A

29

S P SU A S P su 9 w S P su A w S P SU A

2s

Paramysis kroyeri Paramy& nauveli Paramysia ponticu Paramy& ulkkyi

46'N

Praunus Jee.2cosis

58'N

W. Europe

X

SPSUAa

2

55'N 57ON-54'"

Denmark Scotland, Irish Sea

X

Praunus inermis

sp su a S P SU A w

2 2

55"N 56"30" 55"N 56'N-40°N

Denmark Scotland Denmark Atlantic

X

Praunus neglectus Pseudomma aflne

46"N

Black, Azov Seas Roscoff Black, Azov Seas Russia

Rhopalophthalmus brisbanenaia Schistomysis kervilbi Schietomysia ornakc Schistomysis spiritus

27"30'S

Australia

58"N 56'30" 5BoN-54'N

Scotland Scotland Scotland, Irish Sea

Siriellu annata

58ON-54'N

Scotland, Irish Sea

Siriellu clausii SpelaeQmyiri8 longipw

43'N 10"N

France India

X X

X

X

X X

X X X

X X

sp su a sp su a sp su a w (9)

09

a.

29 2s

2-3 2 2

-

S P SU A w

39

SP SU A w SP su a W S P SU A w

2 1 3-4

all year?

-

SPSUAa S P su

2 2(?)

Referencea Boroditch and Havlena (1973) Zatkutskiy (1970) Labat (1957 Zatkutskiy (1970) Boroditch and Havlena (1973) Liao (1951); McLusky and Heard (1971); Maucbline (1971~) Blegved (1922) Liao (1951); Mauchline (1965) Blegved (1922) Mauchline (1971~) Blegvad (1922) Mauchline (1971e); Wigley and Burns (1971) Hodge (1963b) Mauchline (1971d) Mauchline (1970a) Liao (1951); Mauchline (1967) Liao (1951); Mauchline (19710) Macquart-Moulin (1965) Nath (1973)

The frequency of observations is shown as few or seasonal (many). Periods of intense breeding are indicated by capitals: SP(sp), spring; SU(su), summer; A(a), autumn; W(w), winter.

10.

POPULATION DYNAMICS

197

categories of sexual maturity were determined within the populations of many of the spec?es at frequent intervals throughout the year but only occasionally in the case of other species (few observations). Most of the species investigated have extensive breeding seasons while a few do not. The patterns of reproduction and succession of generations vary between different species and between populations of the same species at different latitudes. The life history of any one species, however, is usually recognizable as one of the following types : (a) species requiring more than two years to achieve sexual maturity such species produce less than 0.5 generations per year ; (b) a species that attains sexual maturity and breeds at an age of' two years, such a species produces 0.5 generations per year; (c) a species that breeds once per year, that is, produces one generation per year ; (d) a species producing two generations per year ; (e) a species producing three generations per year ; (f) a species producing more than three generations per year.

Species producing less than 0.5 generations per year. The bathy pelagic Gnathophausia ingens is the only species so far investigated that has a long life history. Childress and Price (1978) show that it attains sexual maturity at an age of over seven years and that the generation time i s eight years, the young being carried within the marsupium by the female for more than a year. The female produces a, single brood in her lifetime, that is she is semelparous, whereas the majority of other mysids are iberoparous, producing more than one brood per female. Population length-frequency histograms have a peculiar form in this life cycle. Childress and Price found them to be polymodal, the individual modes clearly distinct and corresponding to successive instars. The total number of instars in the life history is 13, the first two living within the marsupium of the female. Species producing less than one generation per year. One species, the freshwater Mysis relicta, has been shown to have a two year life cycle in certain regions (Fig. 49). Lasenby and Langford (1972) found that this species required two years t o attain sexual maturity and breed in Char Lake at 74"N latitude, thus providing confirmation of a two year life cycle first proposed for this species in the Great Slave a t 62'N by Larkin (1948). Carpenter et al. (1974) suggest that this species also has a two year life cycle in Lake Superior. Two other species, M . litoralis and M . oculata, may have two year life cycles in the East Siberian and

L 98

THE BIOLOGY OF MYSIDS

Breeding

ds and 8 s Immature

-

PIG.49. Schematic diagram showing the seasonal composition of a population with a two-year life cycle. The arrow indicates production of the next generation by the breeding adults (closest hatching). The juveniles (open hatching) develop into immature adults (closer hatching) and in some populations may mature at an age of 1 year (broken hatching) t o produce a first brood of young.

Laptev Seas (Geiger, 1969); the data, especially on the latter species, are inconclusive and require confirmation. A species that normally matures to breed at an age of two years may mature and breed a t an age of one year in warmer parts of its distribution; individuals of such populations may survive t o an age of two years and breed a second time. This has not so far been demonstrated in a mysid, Mysis relicta having its life span apparently shortened to approximately one year in warmer regions. A peculiar life cycle, first suggested by Fiirst (quoted by Carpenter et al., 1974) for populations of M . relicta in some Swedish lakes, may also exist in this species in Lakes Huron and Ontario. An alternation of generations on an 18 month time scale may take place (Fig. 50) ; breeding females in the summer may produce a generation that matures at an age of 18 months to breed in the winter and produce a generation that matures in 18 months to breed in the summer. This form of life cycle, producing 0.7 generations per year, requires confirmation. A further variation, described by Hakala (1978),may also be another example of alternation of generations in this species. It breeds in Lake Paajarvi, Finland, in

10.

I I Spring I I

Winter

199

POPULATION DYNAMICS

1

Summer

I

I

Autumn

'

Winter

FIG.60. Schematic diagram of the lifecycle of My& relictu in which there is an alternation of generations. Winter breeders produce a winter generation of young that matures 18 months later to breed in the summer; this summer generation of young matures 18 months later to produce a winter generation and so on. Hatching as in Fig. 49.

winter and two cohorts of young are released in the spring. Young released in April and early May mature during the summer to breed the following winter; they die a t an approximate age of 13-14 months. Young released in late May and early June, forming a second cohort, remain immature throughout the first winter but grow rapidly the following summer to breed in their second winter; females of this cohort die a t an approximate age of 22 months and males at 1&18 months. Dr R. M. Laws of the British Antarctic Survey kindly supplied the two samples of Antarctomysis maxima analysed in Fig. 51. The larvae of tihis species probably emerge from the marsupium at a length of 4-5 mm although they have not been observed, the smallest juvenile caught in these samples having a length of 10 mm. Both samples are trimodal and it seems probable that two years are required t o attain sexual maturity and breed. This species attains a body length of over 70mm and so it is conceivable that some two year old animals may survive to an age of three years and breed for a second time. Several of these longer lived mysids may be semelparous like Gnathophusia ingens. Mysis relicta in northern lakes carries the young in the

200

THE BIOLOGY OF MYSIDS

6

10

14

18

Body length ( mrn) 22 26 30

34

38

42

30 January, 1969

FIG.51. Population histograms o f samples of Antarctomysis m a x i m collected b y the British Antarctic Survey. Juveniles and females are shown above the lines, males below the lines. The transverse hatched lines indicate possible correspondence between modes.

marsupium for a t least 150 days at environmental temperatures below 10°C (Table VI, p. 64). Consequently production of a second brood by females is unlikely. The same is probably true in northern populations of other Mysis species and in Boreomysis arctica and B. nobilis. Antarctomysis maxima, on the other hand, may be interoparous, a proportion of females surviving to be three years of age and produce a second brood. Species having these relatively long life cycles are probably of a larger body size than species producing one or more generations per year. These types of life cycles should be looked for in species living in the deep neritic and the mesopelagic and bathypelagic environments. The life cycle shown in Fig. 49 is probably recognizable in the majority of cases by a clear separation of age groups or modes in population

10.

201

POPULATION DYNAMICS

histograms. Histograms of the cycle in Pig. 50 will contain three or four modes more closely spaced than the modes of the cycle in Fig. 49.

Species producing one generation per year. Mysis stenolepis in Passamaquoddy Bay and the Gulf of Maine has a conventional life cycle spanning one year and illustrated schematically in Fig. 52. The

1

I

I

Winter

I

Spring

Summer

I

I

I Autumn

I

Winter

1

FIQ. 52. Schematic diagram showing the seasonal composition of a population of a species producing one generation per year. The solid arrow indicates production of the next generation by the breeding adults. A second brood (broken arrow), lor cvcn a third brood, may he produced by the females during the breeding 8eason. Significance of hatching as in Fig. 49.

young are released from the marsupium during a restricted period in the spring and attain sexual maturity the following spring when th2y breed (Smith, 1879; Amaratunga and Corey, 1975). Females carrying embryos only occur in the population during the period April to June and no breeding takes place a t other times of the year. Populations of M . mixta in the general area of the Gulf of Maine probably have this type of life history (Smith, 1879; Wigley and Burns, 1971) but confirmation is required. Populations of M . relicta can also have a life cycle spanning one year. Lasenby and Langford (1972), investigating populations in Stony Lake (Table XXXIV), found that eggs passed into the marsupium in December and young developed there to be released

202

THE BIOLOQY OF MYSIDS

in May. The breeding adults survive for some time into the summer but no later. The young attain sexual maturity and breed the following winter. This species is also an annual in soutsh-eastern Lake Michigan where Reynolds and De Graeve (1972) found that it breeds continuously throughout the year, although most intensively in the summer. Morgan and Beeton (1978) note that young are recruited into the population near Milwaukee on Lake Michigan a t distinct intervals throughout the year. The population consequentdy consists of cohorts, the females in any one cohort maturing to breed a t an age of one year and a significant proportion of them surviving, moulting and breeding for a second time four months later. Two coasta.1 neritic species in Scotland, Mysidopsis didelphys and Schistomysis ornata, are annuals. The former breeds in the autumn and winter, while populations of the latter species breed in the winter and spring, although some breeding individuals occur during all seasons of the year. These are probably semelparous species. It appears that some populations of an annual species may be semelparous while others may have interoparity. The factors controlling this are not clear. These life cycles extending over a full year probably apply to certain species in the latitudinal bands 40-60" and to neritic rather than littoral species. The breeding females may produce one or more (hatched arrow in Fig. 52) broods during the breeding season.

Species with two generations per year. The most simple structure of this life history is shown in Fig. 53 by the solid arrows. An overwintering generation matures and breeds in the spring to produce a spring generation. The members of the overwintering generation die after breeding. The spring generation grows and matures during the spring and summer to breed in the late summer to early autumn period and produce a new overwintering generation. The members of the spring generation also die immediately after mating. Consequently, except for short periods of time the population consists of only one age group. The simplicity of structure of this life history depends upon all members of each generation maturing and breeding within a short period of about two months. The life history of none of the species so far investigated conforms to this simple structure, although that of Praunus flexuosus in Scotland and Denmark approximates to it (Mauchline, 1971~).In P. jlexuosus, however, the breeding period is long, extending from February to July and involving the production of successive broods by the females, indicated in Fig. 53 by the hatched arrow. Females reared in the laboratory can produce a brood of young every three or four weeks. There is no method a t present for identifying successive

10.

203

POPULATION DYNAMICS

-

--

I

I

I

Winter

I 1

1

1

Spring

I

+-

Summer

I

I

Autumn

Winter

I

FIG. 53. Schematic diagram showing the seasonal composition of a population of a species producing two generations per year. The breeding females may produce more than one brood of young (additional broken arrows). Body size a t sexual maturity is reduced in summer breeders. Significance of hatching as in Fig. 49.

broods in wild populations. Females in a population of Neomysis americana, studied in Passamaquoddy Bay by Pezzack and Corey (1979), produce broods that are identifiable as modes in the lengthfrequency histograms; this is a result of a high degree of synchronous breeding within the population. This pattern of life history is shown in Fig. 54. The overwintering generation matures in the spring and the first brood is released from the marsupium by all females within a short period of time, giving rise to the first cohort of the spring generation.

Winter

I

I

Spring I

Summer

I I

I

Autumn

I

Winter

FIU.54. Schematic diagram showing an alternative seasonal composition of a population producing two generations per year. See text for further explanation. Significance of hatching 8s in Fig. 49.

204

THE BIOLOGY OF MYSIDS

A second brood is then produced by the females and results in a second cohort of the 'spring generation. The breeding adults die soon after producing the second brood. The first cohort matures and breeds before the second cohort so that the new overwintering generation initially consists of two cohorts. These, however, merge in the winter to form a more or less uniform population. The summer generation does not survive far into the winter. The majority of populations of species listed in Table XXXIV as producing two generations per year have extensive breeding seasons so that size modes within the populations are often partially obscured. This situation is further complicated by a decrease in size of the mysids during the summer period. I n the spring generation, the juveniles develop secondary sexual characteristics and the males and females mature to breed a t smaller body sizes relative to individuals of the overwintering generation. This is indicated in Figs 53 and 54. Lack of synchrony in brood production by the females, combined with the production of successive broods by any one female, causes difficulty in deciding whether there are two generations in the year as opposed to three generations. This is further accentuated in latitudes close to 40" where a population might be expected to have an increased number of generations and individuals a shorter life span. Species having two generations per year appear to occur in latitudes higher than 40".

Species with three generations per year. This form of life history is difficult to determine in natural populations and is illustrated in Fig. 55. Species producing three generations per year usually exhibit more or less continuous breeding with periods of seasonal maxima throughout the year. I n addition, any one female will produce one, two or more broods. Consequently, the populations consist of very mixed age groups and individual generations are obscured. This happens in species such as Mysidopsis gibbosa, Neomysis integer and Schistomysis spiritus in Scotland and in other species listed in Table XXXIV. The overwintering generation produces a few young throughout the winter ; this winter breeding may be absent in populations of the species in an area of more severe climatic conditions. Breeding, however, becomes intensive in the spring and the production of the spring generation can normally be recognized. Most females in species with this life cycle probably produce two broods of young because the resulting spring generation often appears to be bimodal. The spring generation matures and breeds in the summer t o produce a summer generation. Breeding females from the overwintering population may survive t o this time, although in small numbers, and still be producing

10.

Winter

I I

205

POPULATION DYNAMICS

I

I

Spring I

Summer ~

I

I Autumn I Winter

~~

FIG. 55. Schematic diagram showing the seasonal composition of a population of a species producing three generations per year. Format of diagram as in Figs 49, 52 and 53.

young. The breeding females of the spring generation may produce two or more broods, but usually die by early autumn. The summer generation grows during the late summer and early autumn, at least its oldest members (Fig. 5 5 ) maturing to breed in the autumn and produce an overwintering population. Members of the summer generation, especially the younger ones, very frequently also form part of the overwintering population, as indicated in Fig. 55. The proportion of the summer generation that overwinters is probably dependent upon the autumnal climatic conditions. The early onset of cold weather in the autumn may increase the proportion that does not mature to breed in the autumn but that survives the winter to breed either during the winter or in the following spring. The schematic representation of this form of life history is simplified in Fig. 55. Continuous breeding within populations means that there is a continuous, although seasonally variable, production of young. Neverthelees, this basic pattern can often be discerned relatively easily in the populations. Detection of the pattern is simplest in populations where winter breeding is minimal and the spring generation is easily recognizable. This situation usually leads to a summer generation that can also be seen as a mode, although less well defined, in the length-frequency histograms. Development of the secondary sexual characteristics as well as the attainment of sexual maturity and production of young takes place at smaller body sizes in members of the spring and summer generations than in members of the overwintering population (Fig. 55).

206

THE BIOLOGY OF MYSIDS

This type of life history is probably most common among shallow living neritic and littoral species occurring in latitudes between about 25" and 50".

Species with more than three generations per year. The presence of more than three generations per year results in extremely complex population histograms (Fig. 56). Populations of Metamysidopsis elongata, probably of Mysidiurn columbiae and in some years Schistomysis spiritus (Table XXXIV) are in this category. Breeding is more or less continuous throughout the year and each female produces several broods. Generations overlap and cannot be distinguished. The simplified

--------1

I

Winter

I

Spring

I

I

Summer

I

1

Autumn' Winter

pattern shown in Fig. 56 is rarely discernible in the population. Its presence is deduced from laboratory observations on the rates of growth, attainment of sexual maturity and brood production. Most species with this life cycle are littoral or shallow neritic and occur in shoals in latitudes lower than 40".

As explained above, the life histories illustrated in Figs 49-56 are idealized and simplified. Most natural populations, but not all, are less regular owing principally to the presence of extended, if not continuous, breeding seasons and the production of more than one brood of young by most females. Summer breeding individuals, however, are usually smaller than individuals of the same species bseeding at cooler times of the year. Consequently, examination of length-frequency histograms of ovigerous females often aids in the interpretation of the histograms of

10.

POPULATION DYNAMICS

207

the whole population. Most of the species listed in Table XXXIV are shallow coastal and littoral species with the addition of a few species occurring in freshwater habitats. The biology of none of the more strictly offshore species, whether epipelagic or deeper living (100-500 m depth) hyperbenthic, has been examined. Their life cycles are probably simplified, span one or two years, and are comparable with those of species such as Mysis relicta, M . stenolepis, Mysihpsis didelphys, Schistomysis ornata and Leptomysis gracilis. Mesopelagic and bathypelagic species, such as the large sized Cnathophausia species, may all have an increased longevity as shown for C. ingens by Childress and Price (1978). Rates of growth of individuals of various species have been computed by the following authors : Gnathophausia ingens by Childress and Price (1978); Hemimysis speluncola and Leptomysis lingvura by Gaudy and Guerin (1979); Metamysidopsis elongata by Clutter and Theilacker (1971) ; Mysis gaspensis juveniles by Dadswell (1975) ; M . relicta by Larkin (1948), Lasenby and Langford (1972), Reynolds and De Graeve (1972) and Hakala (1978) ; M . stenolepis by Amaratunga and Corey (1975) ; Neomysis intermedia by Murano (1964b) ; Praunus species by Blegvad (1922). Growth rates of many of the species can be computed from the data given in the papers quoted in Table XXXIV. Growth of individuals is faster in generations produced in warmer months of the year than in colder months. Consequently, growth rates of a species can vary between generations. The longevity of individuals of a species can also vary markedly between generations. Individuals in a summer generation may have a life span as short as 6-8 weeks, while overwintering individuals of the same population may live for 5-7 months. Growth in body size of a mysid takes place, as in all crustaceans, principally at moulting and so a truly descriptive growth curve of an individual has a stepwise form. A certain amount of growth in body size can, however, take place between moults. Adult female mysids with embryos in the marsupium do not moult although they may, in the case of Cnathophausia ingens, Boreomysis arctica or Mysis relicta (Table VI, p. 64, carry their embryos for several months. Successive stages in the development of the embryo and larvae can be recognized (Chapter 3) and these are equivalent to successive times in the course of the intermoult period. This situation allows an examination of the potential increases in body size that can be accommodated without the necessity of moulting. Measurements were made of the body lengths of females with embryos or larvae at known stages of development within the marsupium (Table XXXV). Females with eyeless larvae

TABLE XXXV. MEAN BODYLENGTHS f (I AND NUMBERSOF FEMALES WITH DIFFERENT LARVAL STAGES IN THE M A R S ~ I U M . THERATIOS OF THESE LENGTHS ARE GIVEN (Mauchline, 1973a) Lengths (mm)of females with Numbers of femalea with “Eggs”

Eyehs (2)

EYd (3)

Species

(1)

Erythrops serrata Leptomysis gracilis Leptmysis linyvura Mysidopsis didelphys Mysidopsis gibbosa Neomysis integer Paramysis arenosa Praunua flexuosus Praunus inermia Praunus neglectus Schistomysia kervillei Schistomysis o d a Schistmysis apim’tua Clyde Loch Etive

6.3 rf: 0.7 12.0 f 1.4 9.4 f 0.7 11.9 f 1.2 4.7 f 0.6 13.5 f 1.5 7.5 f 0.8 19.2 f 2.2 12.6 f 1.1 19.1 f 1.8 10.8 f 1.1 12.2 rf: 0.7

6.6 f 1.0 7.4 f 1.0 12.9 f 1.5 12.4 f 1.4 10.0 0.4 9.8 f 0.9 11.7 f 1.3 12-1f 1.1 4.9 f 0.6 5.1 f 0.6 13.2 f 1.6 13.6 f 1.7 7.9 f 0.8 7.7 f 0-8 19.1 f 2.0 19.7 f 2.0 12.9 & 1.2 2043 1.7 20.1 f 2.2 11.2 f 1.3 11.3 f 1.2 13.9 f 1.2 12.7 f 0.7

11.4 f 0.9 11-7f 0.9

11.8 f 1.0 12.2 f 1.0

12.1 rf: 1.0 12.2 f 1.0

Ratioa of bngths

(1)

(2)

(3)

2/1

3/1

16 327 12 13 283 353 174 250 318

36 462 12 26 316 338 324 342

13 113 5 12 95 158 99 90

1.05 1.03 1.04 0.98 1-04 0.98 1.03 0.99

1-17 1.08 1-06 1-02 1.09 1.01 1.05 1-03

6

203 6

14 415 21

9 156 7

1.05 1.04 1.04

1.09 1.05 1-14

367 250

347 42 1

133 169

1.04 1.04 Mean 1.03

1.06 1.04 1-07

638

1.02

10.

209

POPULATION DYNAMICS

were, on average, 3 yolonger in body length than females carrying early embryos (“eggs”), while their body length had increased by about 7% ab t,he time when the larvae had developed stalked eyes. The abdomen was found to increase in length relative to the carapace during this time as instanced in Fig. 57 for Praunus jlexuosus. The intermoulb period of females carrying young is undoubtedly extended in duration

-

0

I

I

1

I

I

FIG.57. Abdomen length related to carapace length in female Praunusjexuosw carrying “eggs” (dots) and eyed larvae (triangles)in the marsupium (afterMauchline, 1973a).

relative to that of a female of the same body size but not carrying young. Consequently, a 7 % increase in body size between moults probably overestimates the comparable intermoult increases in body sizes of juveniles, males and immature non-breeding females. Most of the increase in body size is associated with moulting, and a model of growth has been developed that applies to decapod and several other types of crustaceans (Mauchline, 1976, 1977b, c). The duration of the intermoult period was found to increase logarithmically while the

210

THE BIOLOGY OF MYSIDS

growth factors, that is the increment in length expressed as a percentage of the premoult length, decreased logarithmically when related to body length or successive moult number. Very few observations of the intermoult periods of mysids have been made, but the relationship appears to exist in the data in Figs 41 and 42 (p. 143). Growth factors, related to body length, can be calculated from the data of Ishikawa and Oshima (1951) and Matsudaira et aE. (1952) and these are plotted logarithmically against carapace length in Fig. 58. The equations of the two regression lines are:

Neomysis japonica

log G F = 1.423 - 0.194 CL

Gastrosaccus vulgaris

log GF = 1.406 - 0.231 CL

where G F is the percentage growth factor and CL is the carapace length. The correlation coefficients of both regressions are significant at the 0.1% level. Gaudy and Guerin (1979) reared Hemimysis speluncola and Leptomysis lingvura and measured increase in length of the exopod of the uropod at successive moults. The percentage growth factors did not decay logarithmically when related to successive moult number (Fig. 59). The animals were reared at the four temperatures Total length (mrn)

40

20

'

" O r b '

100

60

80

I

'

I

I

1

1

1

1

140

120 I

I

I

1

1

1

'

201.

1

t

1

1.0

1

1

1.5

1

1

2.0

1

1

1

1

1

1

1

J

2.5

Carapace length (mm) FIG. 58. Growth factors expressed as the percentage increme in carapace length of Neomysis japonica (dots and Gmtrosaccus vulgaris (triangles) and as percentage increase in total length of Gnathophawia ingens (stars) related to the increasing size of the animals. The equations of the two lines are given in the text. Data from: Ishikawrt and Oshima, 1951; Matsudairs et al, 1952; Childress and Price, 1978.

10.

POPULATION DYNAMICS

21 1

40r 30 -

-8

20 -

s

c

u

0

5 10-3

2

7-

0 0

4-

10°C 14°C 18°C 22°C

u 3 5 1

7

9 Moult number FIG.59. Growth factors, expressed as percentage increase in the length of the exopod of the uropod, at successive moults in Leptomysis lifl,gvura and Hemimysis speluncola at different temperatures. Data from: Gaudy and Guerin, 1979.

shown in Fig. 42 where the information on the corresponding intermoult periods is given. There was much variation in the growth factors and it does not appear to be related to the environmental temperatures. The data for Hemimysis speluncola are slightly more regular than those for Leptomysis lingvura. The only other measurements of growth factors a t successive moults are for the bathypelagic Gnathophausia ingens (Fig. 5 8 ) by Childress and Price (1978). They were obtained from population length frequency histograms which are polymodal, the modes corresponding to successive instars. The larva within the marsupium moults on emergence and has a growth factor of 24.3%. The subsequent seven moults have factors ranging from 1 9 4 % to 20.4%; in other words, the factor remains constant. The factors then decay from 20.4% to 14.9% over the following three moults, the last three moults of the normal life cycle. In both cases, the growth factors for the final series of moults of Hemimysis speluncola and Gnathophuusia ingens decay in an approximately logarithmic manner.

212

THE BIOLOUY OF MYSIDS

Consequently, the available data on the pattern of increase of the durations of the intermoult periods fit the model of growth. The logarithmic decay of growth factors is present in the data for Neomysis japonica and Gastrosaccus vulgaris. The results for Leptomysis lingvura and Hemimysis speluncola are peculiar in the context of crusbacean growth in general; the growth factors increase to a maximum or modal value and then decay (Fig. 59). This pattern of growth factors occurred in Forster’s data (quoted by Mauchline, 1977b)on the growth of Palaemon serratus in culture. A metamorphosis is involved in this animal and it seems possible that the immediately succeeding larval stages were adversely affected by the conditions of culture; the growth factors were depressed but recovered, by increasing at the successive three moults, after which they decayed in a logarithmic manner. Similar effects of the culture procedure may have affected the growth factors of the early moults in the cultures of Gaudy and Guerin and only further observations on these and other species will clarify the situation. The bathypelagic Gnathophausia ingens may be specially adapted to its environment, as discussed by Childress and Price (1978); the growth factors remain constant until just before the animal starts to mature sexually. The model, therefore, described by Mauchline is probably applicable to all mysids with the exception of bathypelagic species, if G. ingena is typical of them. Assuming that the model is appropriate, laboratory observations of growth factors and intermoult periods can be used in an empirical fashion to construct models of growth rates of individuals and populations. These models can aid in the interpretation of the environmental data, especially where the generations are obscured through continuous breeding of the population. The detailed methods of applying the model are fully explained in the companion work on euphausiids in the second part of this volume, where it is used to examine the growth of Euphausia superba Dana. The number of instars in the life of a mysid appears to be markedly fewer than in many other crustaceans. The first instar lives within the marsupium where it moults and the second instar also dwells within the marsupium. This instar moults as it is released, or shortly after release, from the marsupium. There are a further 10 or 11 instars in Gnathophausia ingens before sexual maturity is achieved (Childress and Price, 1978). Neomysis japonica requires 11 post-marsupial instars to attain sexual maturity (Ishikawa and Oshima, 1951). As mentioned previously, summer generations breed at a smaller body size than overwintering generations. Matsudaira et al. (1952) found that the small sized breeding individuals have 11-13 post-marsupial instars, while the

10.

POPULATION DYNAMICS

213

larger sized animals of the winter generation have 10-21 instars. Similar numbers of instars occur in the corresponding generations of Leptomysis lingvura according to the data given by Gaudy and Guerin (1979). Hemimysis speluncola, however, a cavernicolous species, has 10-11 post-marsupial moults. The difference in body size between breeding adults of different generations is less in this species than in Leptomysis Eingvura and consequentdy only one or two extra moults will be required by overwintering individuals. Moulting, and therefore growth in body size, may be inhibited by low winter temperatures in middle and high latitudes. Such inhibition has to be taken into account in analysing growth of individuals in such regions. Variations in size at sexual maturity not only exist within populations of a species a t different seasons but also between populations in different areas at the same season. Wenner et al. ( 1 974) examine size at sexual maturity and the growth rate of crustaceans in general but include mysids as examples. Their model assumes that the rate of growth and the time required to attain sexual maturity are related. Female mysids generally grow larger than males. Clutter and Theilacker (1971)found that immature male and female Metamysidopsis elongata increase in body size a t the same rates up to an age of about 30 days. Thereafter, although males moult more frequently, as shown by the data in Fig. 41, they grow at a slower rate, This means that the rate of decay of the growth factors at successive moults is faster in males than in females. The growth factors of newly liberated juveniles of Gastrosaccus vulgaris and Neomysis japonica are close to 20% in Fig. 58. The latter species attains a maximum carapace length of about 3.2 mm in females and, by extrapolation in Fig. 58, growth factors will have decayed to values of 6.3%. Gastrosaccus vulgaris attains a maximum carapace length of about 4.0 mm in the females and the growth factors at this size will be about 3%. This is also true of the growth factors of Leptomysis Eingvura and Hemimysis speluncola in Fig. 59. It is not true of the bathpelagic Gnathophausia ingens (Fig. 58). Excepting G . ingens, these ranges and rates of decay of the growth factors are comparable with those found in other crustaceans ; immediate post-larval moults usually have growth factors in the range 20-26% and these decay to factors of 3-6% near maximum size, normally attained after breeding. Sex ratios within populations of mysids are variable and females frequently outnumber males. Brattegard (1973, 1974a) found that the ratios in 15 species ranged from 1.1 to 5.5 in favour of females; one species Heteromysis elegans had a ratio of 0.9 females t o 1 male. A ratio of 1 : 1 appears to exist in populations of Mysis relicta when combined samples for the year are considered (Larkin, 1948; Reynolds and De

214

THE BIOLOGY OF MYSIDS

Graeve, 1972). The ratio, however, normally varies between samples and between seasons (Kinne, 1951 ; Zatkutskiy, 1970; Brown and Talbot, 1972; Reynolds and De Graeve, 1972). This is illustrated well in Figs 60 and 61 where the changing sex ratios found in samples of species from the west coast of Scotland are shown. No consistent reasons for the observed fluctuations have so far been found. Mortality of females after breeding appears to be related to a subsequent dominance of males in

3L

Erythrops serrafa

2

Mysidopsis gibbosa

Neomysis integer

r

..

I J l F l M l A l M l J I J l A l S 1 0 " I FIG.60. Sex ratios, expressed as the dominance of females above or of males below the hatched line, of species from the west coast of Scotland in different months of the year. Mysidopsis gibbosa :dots, 1964; triangles, 1965. Neomysis integer: dots, Scotland; triangles the Baltic Sea (Kinne, 1951).

some cases. Hakala (1978) found that males died immediately after copulation in Lake Paajarvi, Finland. Sexual differences in the minimum body lengths at which the secondary sexual characteristics become recognizable may be responsible for other fluctuations. Sexual differences in the longevity of males and females and the larger body size attained by females affect sex ratios in populations dominated by larger size groups. This operates in breeding populations in the spring when, in many cases, only mature adults and juveniles of indeterminable sex are present. The greatest imbalance found in the sex ratio of

10. POPULATION

215

DYNAMICS

a population is 19 : 1 in favour of females in Praunus inermis in November (Fig. 61). Wigley and Burns (1971) observed a ratio of 59 : 1 in favour of females in their collections of Mysis mixta, and suggest that the males may live in a different habitat from the females ; no clear segregation of males and females of a species has yet been demonstrated. A geographical difference in the sex ratio occurred in populations of Eucopia unguiculata (hanseni); Casanova (1977) found that the ratio in favour of females was 1-4 : 1 in the eastern Atlantic off North Africa, 2 : 1 in the Gulf of Gascogne and that a ratio of 1 : 1

Praunus tntvmis

,.A,

J F M A M J J A S 0 N D FIO.61. Sex ratios, expressed as in Fig. 60, of Praunw species from the west coast of

Scotland.

was present in the Mediterranean. Ratios in the oceanic species h t h p h a u s i a gigas and G. zoea were 1 : 1, in G. gracilis 1-5 : 1 but in G. ingens they have been found to vary from 1 : 1 to as high as 7 : 1 (Clarke, 1962). Changes in the sex ratio within populations can result from differential mortality of males and females as mentioned above. Such mortality has been assumed to exist during several investigations. It is always possible that, in some species, the males may have moved to a different sector of the environment during the day and most observations of populations are made during daylight hours. Evidence of

216

TEE BIOLOGY OF MYSIDS

differential mortality is shown in Fig. 61 in populations of Praunus species. The number of males, especially in P . Jlesuosus, decrease markedly during the spring breeding period and again in the late summer and autumn breeding period. These two species are restricted to zones of occurrence among seaweeds and the males were not found in other sectors of the environment. They had apparently mated and died shortly afterwards. This was also found t o happen in populations of Mysis relicta in Lake P&&j&rvi by Hakala (1978). Conclusive estimates of the rates of mortality are difficult to obtain at most stages of the life cycle. Mortality of larvae within the marsupium of eight species of Scottish mysids was estimated by counting the larvae a t successive stages of development at all seasons of the year and determining mean values. This is necessary because of the seasonal changes in brood sizes that occur in these species. The mean number of young per brood can change by as much as 50% in a period as short as four weeks (Fig. 13, p. 47). The development of the larvae within the marsupium takes two to three weeks, depending upon the temperature (Table VI, p. 64). Consequently, the numbers of young in a brood of early embryos and in a brood of eyed larvae, found in females present in the same sample, are not necessarily comparable because the brood size at egg laying may have changed since the brood with eyed larvae was produced. The results of the analysis (Table XXXVI) suggest that mortality of larvae within the marsupium is about 7% up to the eyeless stage and about 10% over the entire period of marsupial development. Amaratunga and Corey ( 1 975) present a graph showing declining numbers of successive larval stages within the marsupium of Nysis stenolepis. This species releases its young from the marsupium any time after the fist marsupial moult, when the appendages become free, and not all larvae are released from the marsupium a t the same time. The rate of mortality during the developmental period up to and including the f i s t marsupial moult is about 13%. This marsupial mortality was ascribed to premature escape or loss of larvae from the marsupium by Mauchline (1973b) but this opinion has had to be revised. Wittmann (1978) demonstrated retrieval of such larvae by the female and adoption of those that had escaped from the marsupium of another female. Mauchline and Webster confirmed these results and consequently marsupial mortality may be a result of death during development rather than of escape. Fager and CluMer (1968), studying the population dynamics of Metamysidqsis elongata, made several assumptions regarding rates of mortality a t different stages of the post marsupial life cycle; they then generated life tables for a population of this species in California. Age

TABLEXXXVI. NUMBERS OF DIFFERENT STAGES OF LARVAE PER BROOD (Mauchline, 1973b) Ratios of

Mean numbers f Q Species

"Eggs"

28.3 f 16.5 Leptomysis gracilis My&ahpsisgibbosa 16.6f 7.6 Neomysis integer 31.6 f 14.2 Paramysis arenosa 14.3f 6.3 Praunwr jlexuoswr 28.2 f 9.6 Praunua inermis 23*9*11.8 Schistomysis kerwilbi 12.6f 6.8 Schistomysis spiritw Clyde 11.3+ 4.9 Loch Ewe 12.4f 4.5

Number of broods examined

EyelRss

EYd

28.2 f 14.5 13-9f 6.3 27.8 f 13.2 12.2f 5-4 25.8 f 9.1 22.3f 11-9 12.7 f 6-5

26.5 k 14.9 13.7+ 6.7 27.6 f 13.1 10-6f 5.4 26.5 f 10.2 20.8f 11.0 124k 7.0

161 135 301 133 205 278 177

340 164 298 267 279 427 355

10.7f 5.3 12.4k 5.4

12*3& 6.4 10.0k 5.0

347 212

313 343

eyehsl"eggs"

eyedl"eggs"

94 71 151 86 81 148 140

0.997 0.837 0.880 0.853 0.915 0-933 1.008

0-936 0.825 0.873 0.741 0.940 0.870 1.016

123 147

0-947 1.000

1.089 0.807

Mean 0.930

0.900

218

THE BIOLOGY OF MYSIDS

specific survival curves were drawn (Clutter and Theilacker, 1971) from estimates of the greatest median and least rates of mortality estimated in the population. Comparable mathematical exercises can be accomplished using various blocks of data referred t o in Table XXXIV. Childress and Price (1978) found that mortality was low in the early post-marsupial instars of Gnathophausia ingens but high in the later instars. They compare this situation with the constanti rate of mortality assumed by Fager and Clutter to be present over the life of Metamysidopsis elongata. No detailed examination of the pattern of mortality over the life history of a coastal species has been made. Mortality of larvae during marsupial development is 10-15% (Table XXXVI). The rate of mortality in later instars probably changes but varies irregularly in time and space. Predators must account for a considerable proportion of mortality in some populations and species and not in others. The.impact of predators may be greater on some instars than on others, through size selection of their prey. Predators are discussed in Chapter 12. The numbers of young per brood are frequently as low as 2-8 (Table 111, p. 48), and so indicative of very low rates of mortality within populations, even if several broods are produced. The number of young per brood increases as the cube of the body length (Fig. 15, p. 56) and an increase in adult body size is usually associated with a longer life history. Consequently, it can be argued that these relationships show mortality is proportional to length of life. The sizes of broods, however, of species of the same body length can vary by one order of magnitude (Fig. 15). Further, as Childress and Price (1978) instance, Metamysidopsis elongata of 5.3 mm body length at maturity produces more than 14 broods containing 5130 eggs so having a fecundity of about 340, while Gnathophausia ingens of 151 mm body length produces 350 larvae in a single brood. The former species has a longevity of just over 200 days, while the latter lives for 2950 days. Consequently, a considerable range of strategies will have been adopted within the Mysidaceae to maintain population numbers. The f i s t estimates of rates of production within mysid populations were made by Blegvad (1922) in Praunus species.' He suggested that single breeding pairs of P. inermis, P. flexuosus and P. neglectus in Denmark could produce 1306, 764 and 588 young respectively. A comparable estimate of production by a breeding pair of P. inermis in Scottish waters was 2450 young (Mauchline, 1965). Estimates of the production of populations of Xchistomysis spiritus in a relatively sheltered bay in the Firth of Clyde and a relatively exposed bay in Loch Ewe in Scotland suggested that production was greater in the

10.POPULATION DYNAMICS

219

exposed situation (Mauchline, 1967). A model of production within a population of Paramysis intermedia has been developed by Miroshnichenko and Vovk (1973). They found a monthly P/B ratio fluctuating between 0.3 and 3.0, being highest in autumn. Hakala (1978) found P/B ratios of 3 . 6 3 - 8 in populations of Mysis relicta in Lake Paajarvi, Finland. Higher ratios could probably be calculated over periods of four to six weeks during production of summer generations of littoral species such as Schistomysis spiritus. Shushkina (1973) estimated daily P/B ratios indirectly in a population of Neomysis mirabilis in the Sea of Japan as 0.13-0.17, while Richards and Riley (1963) noted a ratio of 3.66 in populations of N . americuna in Long Island Sound.

CHAPTER

11

GEOGRAPHICAL DISTRIBUTION The zoogeographical distribution of the majority of species of mysids is either unknown or has not been analysed. Some 65% of the known species live in shallow coastal regions while the remaining 35% live offshore in the deep neritic or oceanic pelagic environments. The oceanic species, about 250 in number, consist of species with restricted distributions as well as the deep living cosmopolitan species that occur in all three oceans. The distributions of about 30 of these species have been described and attempts made t o relate them t o the discontinuities of the oceans. Records of the following species are shown on world maps by the following authors : Archaeomysis species by Murano (1977a); Amblyops abbreviata and several Boreomysis species by Bjmtein and Tchindonova (1958); Caesaromysis hispida by Birstein and Tchindonova (1958) and Murano (1977a);Ceratolepis hamata, Chlaraspidum alatum and various Eucopia species by Fage (1942) and Birstein and Tchindonova (1958); Gnathophausia gigas and G. gracilis by Fage (1941) and Birstein and Tchindonova (1958); G. ingens by Fage (1941)and revised by Pequegnat (1965); Longithorax species by Murano (1976); Lophogaster species by Fage (1942); Katerythrops oceanae by Murano (1976); Meterythrops robusta and Pseudomma truncatum by Birstein and Tchindonova (1958); Mysis relicta by Holmquist (1963b, 1966); Neomysis awatschensis, N . intermedia and N . mercedis by Holmquist (1973); Teraterythrops robusta by Murano (1975). It is possible to collate records from the literature and produce distributional maps for many species. The records, however, are incomplete in most cases as can be seen by examining many of the maps cited above. There are regions of the oceans that have not been sampled comprehensively for mysids. The mysid fauna of a sea area is difficult to sample and a variety of techniques and sampling equipment has to be used (Fig. 18, p. 67). Sampling methods are discussed in Chapter 3. The coastal and littoral nature of the habitats of many species have resulted in unrepresentative sampling of this order during the early historical periods of faunal exploration. Mauchline and

11.

GEOGRAPHIC&

DISTRIBUTION

221

Murano (1977) have shown that over a hundred new species are described in each ten-year period since 1950, owing t o the increased interest in coastal marine science, especially in tropical and subtropical regions. Many coastal regions of South America, Africa, Australasia and Oceania have not been sampled comprehensively and will no doubt yield many new species and extend the known distributions of some of the present species. Consequently, the mapping of xecords of species at the present time is of limited value. It is important, however, to attempt to identify areas for further investigation. There is also value in examining each geographical region, that is areas that approximate to faunal regions. A considerable quantity of information on the mysid fauna of several regions is available. The literature describing the occurrence of mysids is scattered and large. Gordan (1957) produced a classified bibliography of the literature available up to 1955 and this is an accessible paper. A considerable saving of space is effected in the present work by not quoting papers listed by Gordan unless they axe critical to the subject being discussed. She classified the literature according to species and also according to subject. The distributional literature was divided into that describing freshwater and brackish water species and that describing oceanic species and coastal species. A post-1955 bibliography is given in Appendices I1 and 111.The papers are classified according t o genus while the geographical classification is under regions but not divided into oceanic on the one hand and coastal on the other. Extraction of the relevant literature on a species, or a sea ar'ea, is achieved by reference to Gordan (1957) and Appendices I1 and 111. These lists contain the sources of the distributional information given for each species by Mauchline and Murano (1977)but not cited by them. No attempt is made here to produce an authoritative account of the distribution of mysids. Much more information is required before this can be accomplished. It is even difficult to list species occurring within broad geographical regions with any degree of confidence. Such provisional lists can be made by reference to Mauchline and Murano (1977) where each species has been ascribed to one or more of the regions shown in Fig. 62. Further information has become available since the World List was produced, including the discovery of new species (Table 11, p. 14). Some mysids occur in freshwater environments and are of interest in mariculture systems as food for fish. The majority of species, however, are divided into two broad categories. The first consists of species occurring in offshore deep neritic, epipelagic, mesopelagic and bathypelagic environments. The second comprises shallow neritic, coastal, littoral and estuarine species. This latter group

1 20°

160°

160°

1 20°

80°

40°

O0

40°

80°

--60°

5

-1-

I

1200

160°

- --- --

160°

Fro. 62. Map of geographical regions uaed in defining the distribution of the species in Table 11, Appendix I11 end Mauchline and Murano (1977).

11.

223

OEOOBAPHICAL DISTRIBUTION

conhins the majority of species, many of which have, on present information, extremely iocalized distributions. The following discussion attempts to amplify the brief information given by Mauchline and Murano. Comments are made on species occurring in freshwater environments and on the distributions of oceanic species. These are followed by brief discussions of the occurrence of species within several geographical regions.

Species occurring in freshwater There are about 25 species of mysids that occur frequently in freshwater lakes and rivers (Table XXXVII) although most of them TAFILEXXXVII. SPECIESTHAT OCCURFREQUENTLY IN FRESHWATER L m s AND RIVERS Specie8

cfeographical region

Antromysis almyra Antromysis anopblinae Antromysis cenotemis Antromysis peckorum Gangemysis aasimilis Dkrnyds bahirensia Diamysis pengoi Hemimysis anomala Katamysis warpmhowskyi Limnomysie benedini

Surinam Costa Rica Mexico Jamaica India South Europe East Europe East Europe East Europe Europe

My& litoralis Mysia relicta Nanomyeia 8 i W E ? W i S Neomysie awatschensb Neomysie intermedia Neomysie ntercedia Paramy& baeri Paramysis intermedia

Circumpolar, Arctic Circumpolar, Arctic Thailand Japan, China N. America, Kurile Is.,Japan W. N. America East Europe East Europe

Paramysb kessleri Paramyaia lacuatria

East Europe East Europe

Paramysis ullakyi

East Europe

Spelaeomysie c a r d i s o m Taphrmysia b o w n i

Caribbean, Peru Florida, Alabama

Taphrmysis l o u i & w

Texas, Louisiana

Referencee Brattegard (1977) Bowman (19778) Bowman (1977a) Bowman (197710) W. M. Tattersall (1908) Holmquist (1955) Dediu (1966) Dediu (1966) Dediu (1966) Dediu (1966); Weish and Tiirkay (1975); Boroditch (1978) Holmquist (1959b, 1963a) Holmquist (1963a, b, 1966) W. M. Tattersall (1921) Holmquist (1973) Holmquist (1973) Holmquist (1973) Dediu (1966) Dediu (1966); Boroditch (1978) Dediu (1966) Dediu (1966); Boroditch (1978) Dediu (1966); Boroditch (1978) Bowman (1977a) Beck (1977); Bowman (19778) Bowman (1977a)

224

THE BIOLOGY O F MYSIDS

occur in brackish regions as well.,There are also hypogean species, some of which live in freshwater, some in brackish and a few in truly marine situations (Table XXXVIII). Gordan (1957) lists some 300 papers describing the occurrence of fresh and brackish water species. The more recent literature is listed in Appendix 111. The possible function of marginal caves connecting the marine and freshwater habitats is discussed fully by Riedl and Ozretic (1969). TDLE

XxxvIII.

Species

HYPOOEAN MYSIDS LIVINGI N Qeographical region

Aniaomysia vaeaeuri Antromysis cenotensis A . cubanica

Madagascar Yucatan Cuba

A . juberthiei

Cuba

Hemimysia apeluncola Heteromyaoidea w t t i Leptmyaia burgii Spchomyaia bottazzii S. longipes

France Canary Is. France Italy India

S. nuniezi

Cuba

S. olivae S. quinterenaia S. servatua Stygiomysia holthuiai

Mexico Mexico Zanzibar Lesser Antilles, Puerto Rico Italy Jamaica Italy Jugoslavia

S. hydruntina S. major Stygiomysia sp. Troglomysis vjetrenicmsia

CAVES OR WELLS

References Ledoyer (1974) Bowman (1977a) Bacescu and Orghidan ( 1971) Bacescu and Orghidan (1977) Ledoyer (1963, 1965) Calman (1932) Bacescu (1966) Pesce (1976a) Pillai and Mariamma (1964) Bacescu and Orghidan (1971) Bowman (1973) Bowman (1973) Ingle (1972) Bowman (1976) Caroli (1937) Bowman (1976) Pesce (1976b) Stammer (1933)

The four species Anisomysis vasseuri, Hemimysis speluncola and Leptomysis burgii (Table XXXVIII) live in truly marine caves (Ledoyer 1963, 1974; Bacescu, 1966). Heteromysoides cotti is recorded from a, cave some 6 km from the coast but in which the water level fluctuates tidally (Calman, 1932). Two species, the Stygiomysis species described by Pesce (1975b) and Spelaeomysis longipes, occur in wells. The other species in the genera Spelaeomysis and Stygiomysis are confined t o brackish and freshwater caves and wells, have reduced eyes and a body

11.

GEOQRAPHICAL DISTRIBUTION

225

of the general form shown in Fig. 3, p. 16. The remaining species named in Table XXXVIII occur in freshwater caves. The most widely distributed freshwater species is Mysis relicta which occurs in the Great Lakes and other lakes of glaciated North America and in the lakes of northern Europe and Greenland. It also occurs in the brackish environment of the Baltic Sea. Holmquist (1959b, 1962, 1963a, b, 1966)has made a detailed study of the distribution of this species in the context of marine glacial relict faunas whose occurrence is discussed in detail by Segerstrkle (1962).A related species, M . Zitoralis (Table XXXVII), is also circumpolar and occurs in freshwater lakes although it more commonly occurs in brackish or marine environments. Four other species of this genus, all closely related to Mysis relicta, occur in the Caspian Sea, as discussed later. The rivers of eastern Europe, for instance the Dniestr, Pruth and Volga, that flow into the Black and Caspian Seas contain several species living in truly freshwater environments. The species, listed in Table XXXVII, are Diamysis bahirensis, D. pengoi, Limnomysis benedini, Hemimysis a n m a l a , Katarnysis warpachowskyi, Paramysis baeri, P. intermedia, P. kessleri, P. lacustris and P . ullskyi. Several of these species have been introduced as food organisms for fish in various eastern European reservoirs and their original detailed distributions are becoming difficult to determine (Mordukhai-Boltovskoi, 1979). A recent study of the routes of colonization of the freshwater habitats of this region by Limnomysis benedini has been made by Weish and Tiirkay (1975). Populations of all these species are also present w i t h the Black and Caspian Seas as described below. Most of the remaining species occurring in fresh water (Table XXXVII) have marine coastal distributions. They are listed because they have been recorded on a t least some occasions from truly freshwater situations. Other species such as Mesopodopsis orientalis and Tenagomysis chiltoni in India, M . slabberi and Neomysis integer in Europe have also been recorded from freshwater situations but usually there are obvious routes of carriage from adjacent marine habitats. Neomysis integer is often present in freshwater streams a t low tide but its environment becomes brackish at high tide; other estuarine species also undergo these fluctuations. It has been recorded, however, from freshwater lochs immediately adjacent t o the sea in western and northern Scotland (Scott, 1891a, 1894) but there are no recent records of such occurrences. Further investigation of the distribution of estuarine species and of freshwater habitats adjacent to the coast may increase the list of species with resident populations in truly freshwater environments.

226

THE BIOLOGY OF MYSIDS

Oceanic species These species live in the offshore deep neritic, epipelagic, mesopelagic and bathypelagic environments. Some of those listed by Mauchline and Murano as bathypelagic and others, for example many species in the genus Hansenomysis, ascribed a depth of occurrence may be hyperbenthic, living in close association with the sediments. There are relatively few cosmopolitan species occurring in the Atlantic, Pacific and Indian Oceans. Examples are : Anchialina typica Eucopia sculpticauda Arachnomysis leukartii Eucopia unguiculata Caesaromysis hispida Gnathophausiagigm Echinomysis chuni Cnathophausia gracilis Euchaetomera glyphidophthalmica Gnathophausia ingens Euchaetomera tenuis Gnathophausia zoea Euchaetomera typica Katerythrops oceanae Euchaetomeropsis merolepis Meterythrops picta Eucopia australis Petalophthalmus armiger Eucopia grimaldii Siriella thompsonii These are all meso- and bathypelagic species with the exception of Anchialina typicu and Siriella thompsonii, which are both epipelagic in habit. There are about 12 species that live in both the Atlantic and Pacific Oceans but have not so far been recorded from the Indian Ocean. These include five species of the genus Boreomysis. It is surprising. that at least some species in this genus have not been recorded as cosmopolitan because only one of the 35 species in it, B. insolita, is neither meso- nor bathypelagic in distribution. There are some 30 species living in the Pacific and Indian Oceans but about half of these are shallow living, occurring in at least the western Pacific and eastern Indian Ocean. There are, in addition to these offshore species, a further group of coastal and littoral species occurring across the Australasian region and living in both oceans. Only four species have been recorded exclusively from the Atlantic and Indian Oceans; they are Boreomysis illigi , Euchaetomera intermedia, Longithorax capemis and Lophogaster typicw. The remaining offshore species are confined, on present information, t o one ocean. There are about 70 species restricted t o the Atlantic, about 40 of which occur only in the North Atlantic, north of about 40"N. Some 110 species occur only in the Pacific, 60 of them recorded exclusively from the western Pacific, principally through the work of Ii and Murano. The Indian Ocean has 11 species that have not been recorded elsewhere and there are a few species endemic to i;he Red Sea.

11.

OEOORAPHICAL DISTRIBUTION

227

A detailed examination of the regions of occurrence of several species shown by Mauchline and Murano (1977), raises a number of questions. Parapseudomma calloplura occurs in the north-eastern Atlantic and Mediterranean but has also been recorded by Murano off Japan. Further sampling, and in some cases taxonomic revision of species, may alter the known distributions of many offshore species quite markedly. For example, the genus Hansenomysis consists at present of 15 species, all oceanic, and mostly living in the bathypelagic or hadal environments. The Peru Trench contains seven species, the KurileKamchatka Trench three species, the Antarctic three species and one from the equaeorial Pacific (Mauchline and Murano, 1977). They rarely occur in samples, probably because they are hyperbenthic a%depths of 2000m or more; the Antarctic species of this genus live at much shallower depths, 100-500 m. More frequent sampling at depths greater than 2000 m with a variety of samplers may show that these species are much more widely distributed than is indicated a t present. Biogeographic studies of the distributions of mysids have been neglected and little of value can be gained by extending the present discussion. Much further information is required, as will become clear during the following examination of the mysid faunas of the various oceanic and coastal regions shown in Fig. 62, p. 222. The boundaries of the regions are only approximate and many species cross them in coastal regions. These boundaries, however, are used in Appendix I11 to classify the literature on the distribution of the species. Arctic Ocean Few species occur in the Arctic Ocean, Geiger (1969) and Shih and Laubitz (1978) recording the following:

Boreomysis nobilis Michthyops theeli Mysis litoralis Mysis oculata

Mysis polaris Pseudomma frigidurn Parerythrops spectabilis Meterythrops robusta

There are, however, a much larger numbers of species living south of latitude 60"N, some of which intrude into the Arctic Ocean regularly or sporadically. Some of these have colonized coastal or offshore regions of northern Greenland, northern Europe and northern Alaska and Canada. Such species are:

Acanthomysis pseudomacropsis Amblyops abbreviata Amblyopsoides ohlinii

Boreomysis arctica Boreomysis inermis Erythrops abyssorum

228

THE BIOLOGY OF MYSIDS

Erythrops erythroph#halmu Erythrops glacialis Meterythrops robusta Mysis mixta Mysis relicta

Neomysis intermedia Neomysis rayii Pseudomma truncatum Stilomysis grandis

Sampling for mysids north of 60"Nin both the Atlantic and Pacific Oceans has been sporadic. Little is known about the biology of species living in these high latitudes. Several of the southern guests to this region may be expatriate and their breeding inhibited by the prevailing temperature regimes.

North Atlantic, between about 60"N and 40"N The most comprehensive accounts of the species occurring in this region are those of W. Tattersall (1951) and Tattersall and Tattersall (1951). Other papers are listed in Appendix 111. The species are too numerous to list, being more than 100, some 40 of which are restricted to this region. Faunal lists can be made from the publications of the Tattersalls and from Mauchline and Murano (1977). There are faunal differences between the eastern and western coastal areas of this region. The following species are confined to the western Atlantic :

Boreomysis nobilis Meterythrops robusta ,Mysidetes farrani Mysis gaspensis

Mysis stenolepis Neomysis americana Pseudomma truncatum Stilomysis grandis

Several species such as Mysidopsis bigelowi can occur as southern guests in the coastal regions of the northeastern United States and of Canada ; there are many species endemic to the middle latitudes of the western Atlantic. The 40"N latitude appears to be an approximate faunal boundary for mysids in the western Atlantic but the corresponding boundary in the eastern Atlantic appears t o be south of the entrance to the Mediterranean, a t about 30"N. I n the eastern Atlantic, coastal areas are apparently richer in species than comparable areas in the western Atlantic but this may be an artifact arising from the more comprehensive sampling of workers such as the Tattersalls. There are about 50 species that are restricted to the eastern Atlantic, some 30 of these having distributions that extend southwards into the Mediterranean. Information on the occurrence of these species is given by Tattersall and Tattersall (1951) and Mauchline and Murano (1977). Most prominent among these endemic species are those of Praunus, Paramysis and Schistomysis.

11.

GEOGRAPHICAL DISTRIBUTION

229

All species in these genera are restricted to this area, the Baltic, Mediterranean, Black and Caspian Seas. Some species occur on both Eides of the northern Atlantic but most of these are offshore species, such as Amblyops abbreviata, A. kempi, Boreomysis arctica, B. microps, B. tridens, Erythrops abyssorum, E . erythrophthalma, E. microps, Eucopia grimaldii, Cnathophausia gigas, C.zoea, Parerythrops obesa. There are, however, also the following that occur in coastal regions:

Heteromysis formosa Mysis litoralis Mysis mixta

Mysis oculata Pseudomma afine Pseudomma roseum

A further European species, Praunus jlexuosus, occurs in the Bay of Fundy and the coasts of Nova Scotia. This species was introduced to this coastline from Europe and is now spreading rapidly (Wigley, 1963; Wigley and Burns, 1971). The Baltic Sea The Baltic is a large, shallow, predominantly brackish water appendage of the north-eastern Atlantic. Its western areas are penetrated by several euryhaline species, but in relatively smell numbers compared with their incidence in the adjacent North Sea. These immiqant species are :

Erythrops elegans Hemimysis lamornae Leptomysis gracilis Mysidopsis gibbosa Praunus inermis

Praunus neglectus Praunus jlexuosus Schistomysis ornata Schistomysis spiritus

The dominant resident species within the western Baltic, according to Hesthagen (1973) and Boysen (1976), are:

Qmtrosaccus spinif er Mesopodopsis slabberi

Mysis mixta Neomysis integer

Of these, only Mysis rnixta and Neomysis integer penetrate the northern and inner regions of the Baltic and there they occur along with Mysis relicta. Ackefors (1969) states that M . mixta is common throughout the Baltic proper and especially in the deeper areas. Mysis relicta is a dominant organism among the hyperfauna of the northern regions of Bothnian Bay, while M . mixta occurs further south in the Quark (Brtgge and Ilus, 1973). Ackefors, quoting earlier workers, states that

230

THE BIOLOUY OF MYSIDS

M . relicta is caught i~ Bothnian Bay at depths between 70 and 150 m and may be pelagic by both day and night. It does not occur in areas where salinities are greater than 12%,. One other species, Praunus inermis, has been recorded from the Gulf of Riga and the entrance to Bothnian Bay (Zmudzinski, 1962). The Caspian Sea species, Paramysis lacustris, used as a food organism in the farming of fish, is reported by Zhuravel(l972) to have entered the Baltic via the Newman River in Lithuania into which it had been transplanted from the Caspian. This species may now be in the process of becoming established within the Baltic where the conditions will favour it. AtZantic Ocean, 40"N to 40"s This vast region can be divided into several areas. Starting in the western Atlantic, the first approximat-e faunal area extends from about 40"N off t,he American coastline southwards into the Gulf of Mexico. Tattersall (1951) and Wigley and Burns (1971) describe the species occurring in it. A second area encompasses the Caribbean and possibly northern regions of the South American continent. It is very rich in species as detailed in papers by Bacescu, Bowman and Brattegard. Conversely, sampling, especially in the Caribbean coastal regions, has been extensive and Brattegard has described some 30 new species endemic to this area and the adjacent coasts of Florida, Mexico, Panama and Colombia. A further faunal area possibly exists on the eastern coastline of South America, from about the equator southwards to 40"s. Almeida Prado details the results of sampling a small area of the Brazilian coast, as does Silva. The species recorded are few; one, Metamysidopsis elongata attantica is a form of the eastern Pacific species M . elongata. Much further sampling off this coastline is required before the affinities of its mysid fauna can be discussed. I n the eastern Atlantic, as stated earlier, the northern fauna intrudes southwards to the entrance to the Mediterranean and probably to about 30"N on the African coastline. The mysids of the western African coastline have probably been incompletely sampled. Information for certain areas is available in papers by Bacescu, 0. S. Tattersall and in Grindley and Penrith ( 1 965) and Wooldridge ( 1977). As in the case of the South American coastline, much further sampling is required. The western and eastern parts of this region each contain over 100 species. About 70 of the species recorded in the western area do not occur in the eastern area while some 40 of the eastern species have not been recorded in the west. This probably reflects the variety of

11.

QEOQRAPHIOAL DISTRLBUTION

231

habitats available in the western area but it may also have resulted from more comprehensive sampling of parts of the western area, e.g. the Caribbean. Lists of species can be drawn up for local areas by reference to the literature in Appendix I11 and by recourse to the papers cited above.

The Mediterranean Sea The mysid fauna of the Mediterranean consists of endemic species and species that also occur in the north-eastern Atlantic. A list of more than 30 Atlantic species can be made, by reference to Mauchline and Murano (1977), that also occur in the western Mediterranean; these species, however, do not in most cases penetrate the eastern Mediterranean. The following species, on the other hand, occur in the northeastern AtIantic and throughout the length of the Mediterranean :

Qastrosaccus normani Gastrosaccus sanctus Gastrosaccw spinifer Hemimysis lamornae

Mesopohpsis slabberi Siriella clausii Siriella jaltensis

There are more than 20 species endemic to the Mediterranean.

Anchialina sanzoi, Kainomatomysis foxi and Schistomysis assimilis are only known from the eastern Mediterranean and the other species are :

Anchialina oculala Calyptomma puritani Erythrops peterdohrni Euchaetomera richardi Gmtrosaccus magnilobatus Gastrosaccus mediterraneus Haplostylus bacescui Heteromysis eideri Heteromysis lybiana Leptomysis burgii

Leptomysis peresi Mysideis parva Neoheteromysis mf.klleri Paramysis festae Paramysis helleri Parerythrops lobiancoi Pseudomma chattoni Rhopalophthalmus mediterraneus Siriella adriatica

The Black Sea Paramysis proconnesia appears to be endemic to the Bosphorus. Salinity in the Black Sea ranges, on average, from about 18 to 23%,. Nine Mediterranean species penetrate the Black Sea. Diamysis bahirensis Qastrosaccus normani

Gastrosaccus sanctus Qmtrosaccusspinifer

232

THE BIOLOGY OF MYSIDS

Hemimysis lamornae Leptomysis lingvura Mesopodopsis slabberi

Siriella clausii Siriella jaltensis

All these species, with the exception of Diamysis bahirensis, are widely distributed in the coastal regions of Europe, occurring throughout the Mediterranean, the adjacent region of the Atlantic northwards t o British waters, including the North Sea and, with the possible exception of Leptomysis lingvura, the Baltic Sea. The following six species are endemic t o the Black Sea although Diamysis pengoi also occurs in the rivers flowing into the Black Sea, as mentioned previously :

Diamysis pengoi Hemimysis serrata Paramysis agigensis

Paramysis kroyeri Paramysis kosswigi Paramysis pontica

The Black Sea also shares the following species with the Caspian Sea : Acanthomysis strauchi Paramysis intermedia Hemimysis anomala Paramysis lacustris Katamysis warpachowskyi Paramysis ullskyii Limnomysis benedini

The Caspian Sea The seven species listed above occur in both the Caspian and Black Seas. There are, however, an additional 15 endemic species living in this brackish environment with an average salinity of 13%,. Acanthomysis grimmi Acanthomysis inflata Caspiomysis knipowitschi Diamysis pusilla Paramysis baeri Paramysis bakuensis Paramysis incerta Paramysis kessleri

Paramysis loxolepis Paramysis eurylepis Schistomysis elegans Mysis amblyops Mysis caspia Mysis mawolepis Mysis microphthalma

The four Mysis species are closely related t o the freshwater Mysis relicta. Maximum depths in the Caspian Sea are about 950m. Mysis amblyops, M . microphthalmu and Paramysis loxolepie have bathymetric ranges of about 200-900 m. Mysis cuspia and M . mawolepis occupy the 100-400 m bathymetric zone while the other species inhabit the shallower coastal and littoral areas.

11.

OEOGBAPHICAL DISTRIBUTION

233

The Pacific Ocean The earlier distributional literature on Pacific species is listed by Gordan (1957). The most valuable modern works are those of Bacescu, Birstein and Tchindonova, Hodge, Ii, Murano and Wing. The other relevant literature is listed in Appendix 111. Large areas of the Pacific have not been effectively sampled for mysids. This is especially true of the littoral and other shallow environments of the eastern Pacific stlin lines, of Ocean& and the Aushrahsian region. This is reaected i n the following brief description.

North Pacijic, north of about 4O"N The following species were recorded in the Chukchi Sea by Wing and Barr (1977): Acanthomysis pseudomacropsis Acanthomysis sp. Mysis litoralis Mysis oculata

Mysis polaris Neomysis rayii Pseudomma truncatum

Species occurring in the Bering Sea, the Aleutians and southwards to about 40"N number about 60. Knowledge of the distribution of many of these species is incomplete because the littoral habitats of the coasts of Aleutians and western Bering Sea have not been investigated in detail. Ii (1964a) lists nine species occurring in the Sea of Okhotsk, ten species in the Sea of Japan. A list of some 20 species recorded from the Kurile-Kamchatka Trench and the region to the south and west of it, but not found in the north-eastern Pacific, can be obtained from Mauchline and Murano (1977).

The Pacific Ocean between about 40"N and 40"s The north-western part of this region has been investigated in considerable detail by Ii and Murano. There are about 140 species recorded from the region of Japan, the Philippines and the East and South China Seas. These are fully documented by Ii (1964a) and in the papers of Murano. I n the eastern part of this region, the following species, several of them littoral in habit, have only been found on or close to the American coastline : Acanthomysis costata Acanthomysis macropsis

Acanthomysis sculpta Arachnomysis megalops

234

~

THE BIOLOGY OF MYSIDS

E'uchaetornera plebeja Gnathophausia scapularis Heteroerythrops purpura Heteromysis panamaensis Metamysidopsis elongata Metamysidopsis paci$ca

Mysidopsis californica Neomysis mercedis Scolamblyops oculospinum Siriella pacijica Siriella panamensis Siriella roosevelti

Several species have been found by Bacescu (1967, 19718, b) in the hadal environment of the Peru Trench but have not been recorded from elsewhere as yet. They are:

Amblyops ewingi Hansenomysis birsteini Hansenomysis chini Hansenomysis lucifugus Hansenomysis menziesi Hansenomysis peruvianus

Hansenomysis spenceri Hansenomysis tattersallae Iiansenomysis tropicalis Mysidetes peruana Mysimenxies hadalis

Mauchline and Murano (1977) list some 16 species recorded from the central areas of the Pacific-Hawaii, the Gilbert Islands and other regions of Oceania. Further sampling of the shallow environments of these regions will probably produce many more, and probably new, species. Similarly, many coastal regions of Australia have not been investigated although some 40 species are at present recorded from its coastlines. The mysid fauna of New Zealand is dominated by species of the genus Tenagomysis, although others are also present. The study of occurrence and distribution of mysids throughout the Pacific Ocean is a t a very early stage and systematic investigations over bhe greater part of the Ocean are required.

The Indian Ocean north of about 40"N Regions of the coasts of East Africa and Madagascar have been investigated by Nouvel and Bacescu while Pillai provides much information on the Indian species. The distribution of species within the Indian Ocean is not well known and many littoral areas, off mainlands and islands, have been poorly, if at all, investigated. Faunal lists can be made for several areas but how representative they are of the actual list of species present in these areas is questionable. Much more systematic sampling is required over the greater part of the region with particular attention to inshore environments.

11.

QEOGRAPIIICAL DISTRIBUTION

235

The Red Sea There axe some 20 species that intrude from the Indian Ocean into the Red Sea. These can be obtained from Mauchline and Murano (1977). In addition, the following species appear to be endemic to the Red Sea : Anisomysis levi Paralophogaster i n t e r m d i m Gastrosaccus erythraeus Paralophogaster macrops Gymnerythrops microps Paralophogaster microps Heteromysis digitata Paralophogaster sanzoi Heteromysis gerlachei Pseudan'chialina erythraea Heteromysis kossmunni Siriella dolljksi Idiomysis tsurnamali Siriella serrata Kainomatomysis schieckei Antarctic Ocean Tattersall (1955) describes the species of the Antarctic in considerable detail. She lists eleven species as occurring in the Straits of Magellan; a further 22 occur between 40"s and 60"s (Mauchline and Murano, 1977). The following species penetrate or live permanently in latitudes higher than SO'S : Amblyops antarctica Mysidetes brachytepis Amblyops tuttersalli Mysidetes dimorpha Amblyopsoides crozetii Mpidetes hanseni Antarctomysis maxima Nysidetes microps Antarctomysis ohlinii Mysidetes p o s t h n Boreomysis brucei Paramblyops brevirostris Boreomysis inermis Pseudomma antarcticum Dactylamblyops hodgsoni Pseudomma armatum Eucopia australis Pseudomma belgicae Hansenomysis angusticauda Pseudomma sarsii Hansenomysis antarctica Pseudomma schollaertensis Mysidetes antarctica The above brief account is sufficient to emphasize the lack of information on the geographical faunas of mysids. All faunas are very poorly known with the exception of those of the north-east and north-west Atlantic, Caribbean, Mediterranean and Japanese regions. Many species occur in shallow areas of one to a few metres dept,h. These areas are too shallow to be included within an offshore sampling programme and too deep to be sampled from the shore. This habitat is normally sampled by towing small 0.5 rn diameter nets from boats with outboard engines.

CHAPTER 12

PREDATORS A N D PARASITES Mysids occur widely throughout the marine environment but the most marked concentrations undoubtedly occur in coastal regions. They are, therefore, a potential food resource for coastal fish. This must be especially true of the species of fish that live among the rocks and algal beds in shallow regions where many species of mysids form dense concentrations. Many of these species of fish, however, are of little economic interest and consequently, in many cases, their feeding habits are unknown. Many commercially exploited species of fish migrate into these shallower regions to breed, or for other purposes, and while present there, may feed on the mysids. Several mysids, such as some of those in the genera Leptomysis, Mysidopsis, Neomysis and Schistomysis are known t o form shoals in areas of the continental shelves and are' consequently a potential food resource for a wider range of predatory fish. An epipelagic species such its Pseudanchialina pusilla, which is common in the tropical and subtropical regions of the Indian Ocean, may be preyed upon by the planktivorous fish. Unfortunately, mysids bear a certain resemblance to euphausiids which normally outnumber them in offshore pelagic samples. The mysids tend to be over-looked in such samples and this is probably even more true when examining the stomach contents of pelagic fish. This, in part, may account for the present lack of information on the predation of mysids by planktivorous fish. Miller (1970) found that all age groups of the flathead sole preyed to a considerable extent upon Neomysis kadiakensis but this situation seems to be exceptional among the species of fish so far investigated. The size of prey organism eaten by a fish is often related t o the body size of the fish. Tyler (1972) found that Mysis stenolepis was consumed more frequently by white hake of body length less than 26 cm than by larger fish. Young herring fed on a variety of mysids whereas the adults only ate them occasionally (Hardy, 1924). The olive rockfish studied by Hobson and Chess (1976) preyed most heavily on mysids during the late juvenile period of its life history. Further, many fish perform an ontogenetic migration, the young developing in coastal

12.

PREDATORS AND PARASITES

237

environments while the adults live in more offshore habitats. The European plaice is an example. It has nursery grounds located in sandy coastal bays that are generally inhabited by an assemblage of mysids, often dominated by species in the genera Schistomysis and Paramysis. Edwards and Steele (1968), found that 0-group plaice in one such nursery area of western Scotland consumed Schistomysis spiritus but their diet changed as they developed. This situation could result in predation pressure on the mysids changing seasonally in middle and high latitudes. Moore and Moore (1976) examined the basis of food selection by the flounder and concluded that it depended upon several factors. These included the spatial distribution of the prey, its degree of concealment and its ability to escape and also a conditioning of the fish for certain foods. These factors introduce a further set of variables influencing predation pressure on an individual species of mysid. Moore and Moore concluded that Neomysis integer and Praunus jlexuosus would be quite easily caught by the flounders. The latter species of mysid remains stationary on the surface of the sediment for considerable periods of time and its colouration, or lack of it, may protect it from detection by the fish. The!importance of cryptic coloTation 'in mysids has already been discussed. Neomysis integer occurs frequently in swarms and as Clutter (1969) has pointed out, swarming probably confers protection from predation. This appears to be confirmed by the observations of Hobson and Chess (1978) on swarms of mysids associated with coral reefs. They were surprised that only token numbers of mysids, that occurred in high densities in the water column over the reefs during the day, occurred in the stomachs of planktivorous fish. The information on predation of mysids is widely scattered, and difficult to locate because, of course, the papers are not predominantly about mysids but about one or more of their predators. Consequently, the following account can only be taken as an indication of the widespread importance of mysids in the marine food chains, especially those of coastal regions. Freshwater species of mysids are often introduced into lakes as food for the resident populations of fish, a topic that will be discussed later. Mysis reZictu forms a major component of the diet of several freshwater fish, as detailed in the works of Langford (1938b), Hacker (1956), Rawson (1961), Wells and Beeton (1963), Sparrow et aZ. (1964), Dryer et al. (1965), Grimas and Nilsson (1965), Morswell and Norden (1968), Just (1970), Westin (1970), Northcote (1972, 1973), Reynolds and De Graeve (1972), Nikolaev et al. (1973), Johnson (1975) and Savolainen (1975). Another freshwater mysid, Tenagomysis nowaezealundiae, is

238

THE BIOLOGY O F MYSIDS

eaten by the European perch, Percafluviatilis L., in the Selwyn River, New Zealand (Griffiths, 1976). Epipelagic marine mysids have not been identified among the stomach contents of whales although, as mentioned above, they may have been overlooked among the much more dominant euphausiids that are similar in body form and size. It is, however, more probable that the swarms formed by the mysids are much too small and widely scattered to merit attention by the whales. Nemoto (1959) recorded a young Gnathophausia gigas among the stomach contents of a fin whale caught in the north-eastern region of the Aleutians. Seals, however, apparently feed on mysids to some extent and they have a wider range of swarming species accessible t o them. McLaren (1958) and Dunbar (1941, 1942) found that Mysis oculata is an important element of the diet of the Arctic ringed seal, Phoca hispida Schreber; M . mixta was eaten occasionally. Sergeant (1973) and Dunbar (1942) noted that the harp seal, Pagophilus groenlandicus Erxleben, feeds to a small extent on Mysis oculata and M . mixta. In the Antarctic, the Weddell seal consumes Antarctornysis maxima and A . ohlinii but to what extent remains unknown (0.S. Tattersall, 1965b). Mysids have been recorded in the diets of many species of marine fish. These, however, fall into two broad categories, namely offshore and coastal species. Information on the predation of fish upon offshore species of mysids is lacking. Oceanic species, including bathy-pelagic species, appear to form a minor component of the diet of some planktivorous and deeper living fish. Merrett and Roe (1974), examining the diets of a few mesopelagic species of fish, recorded one unidentified mysid in the stomach contents of the myctophid Lampanyctw cuprarius Tbning as did Clarke (1978) in the diet of L. nobizis off Hawaii. Dragovich (1969) found that the albacore, Thunnus ( G e r m ) alalunga Bonnaterre and the blackfin tuna, Thunnus atlanticw (Lesson), occasionally ate Gnathophausia ingens. I n a later paper (1971) he recorded mysids in the stomachs of yellowfin tuna, ThunPzus (Neothunnus) albacares (Bonnaterre), and the skipjack, Euthynnus (Katsuwonus) pelamis (L.). Macrourid fish are also known to prey upon mysids. Haedrich and Henderson (1 974) found Boreomysis species present among the stomach contents of the benthic rattail Nematonurus armatus (Hector) in the western Atlantic while Pearcy and Ambler (1974) recorded small mysids and Gnathophausia gigas in the food of this species and 'of Chalinura leptolepis (Gunther)iand Coryphaenoides $lifer (Gilbert) in the eastern Pacific. A more recent study by McLellan (1977) lists eleven macrourid fish that include mysids in their diets. Sedberry and Musick (1978) examined the stomach contents of several species of

12.

PREDATORS AND PARASITES

239

fish of the continental slope and rise off the Atlantic coast of the United States. Michthyops parva was eaten by the notacanthid Hatosauropsis macrochir (Giinther), Boreomysis species by the halosaur species of the genus Aldrovandia Goode and Bean, and Boreomysis arctica, B . tridens and Pseudommu roseum by the gadoid longfin hake, Phycis chesteri Goode and Bean. So far, the fish appear to be relatively minor predators of the oceanic mysids but further information is required. Much more information is available about predation of coastal mysids by fish. The most convenient way of examining such records is on a geographical basis because this emphasizes areas and regions where further information is required. The regions used are those shown in Fig. 62. The nomenclature followed for species of fish is that of Bailey et al. (1970) for American species and Hureau and Monod (1973) for north-eastern Atlantic and Mediterranean species.

Arctic Ocean Little information is available on the predation of Arctic mysids by fish. Ponomarenko (1976, 1978) did not identify the species but found that up to 36% by weight of the stomach contents of 0-group cod in the Barents Sea consisted ,of mysids. Herring around Iceland occasionally eat Mysis mixta according to Jespersen (1932).

North Atlantic, between about 60"N and BOON. The following fish of the north-eastern Atlantic include the named mysids in their diet. The list was compiled from the following : T. Scott (19OOa, 1902b), Ray (1914), Todd (1914), Paulsen (1918), Lebour (1919, 1921), Anonymous (1920), A. Scott (1922), Hardy (1924), Savage (1931), Hartley (1940), Redecke (1941), Liao (1951), Tattersall and Tattersall (1951), Jones (1954), Nagabhushanam (1965), Edwards and Steele (1968), Deniel (1974), Kislalioglu and Gibson (1976a), Lande (1976) and Moore and Moore (1976).

Squalus acanthias L., spur dog: Gastrosaccus spinifer. Raja (Raja) brachyura Lafont, blonde ray : Gastrosaccus spinifer. Raja (Raja) clavata L., thornback ray : Gastrosaccus spinifer, other species. Raja (Leucoraja)fullonica I+, shagreen ray: GastrosacczLs spinifer. Raja (Raja) montagui Fowler, spotted ray: Gastrosaccus spinifer. Raja (Amblyraja)radiata Donovan, stamy ray : Boreomysis tridens. Raja (Leucoraja) naevus Miiller and Henle, cuckoo ray : mysids. Raja (Leucoraja) circularis Couch, sandy ray : Erythrops sp.

240

THE BIOLOGY OF MYSIDS

Clupea harengus harengus L., herring : Gastrosaccus sanctus, G. spinifer, Leptomysis gracilis, Mesopodopsis slabberi, Mysis mixta, Neomysis integer, Praunus Jle.xuosus, P. inermis, P. neglectus, Schistomysis ornata, 8. spiritus, Siriella clausii. Sprattus sprattus sprattus (L.), sprat : Mesopodopsis slabberi, M'ysis species, Neomysis integer. Salmo salar L., salmon: Mysis oculata. Salmo trutta trutta L., trout : Neomysis integer. Osmerus eperlanus (L.), smelt : Mysis relicta, Neomysis integer. Syagnathus acus L., great pipefish: young mysids, Neomysis integer. Entelurus aequoreus (L.), snake pipefish : young mysids. Gasterosteus aculeatus L., three spined stickleback : mysids. Spinachia spinachia (L.), fifteen spined stickleback : Neomysis integer, Praunus jlexuosus . Gadus morhua morhm L., cod : Boreomysis arctica, Gastrosaccus species, Mysis rnixta, Jf. oculata, Neornysis integer, Praunus jlexuosus, P. inermis, Schistomysis ornata, S . spiritus, Siriella armata, S . jaltensis, other species. Melanogrammus aeglejnus (L.), haddock : Boreomysis arctica, Erythrops serrata, Gastrosaccus spinifer. Merlangus merlangus merlangus L., whiting : Erythrops elegans, E . erythrophthalma, Gastrosaccus spinifer, Heteromysis formosa, Leptom ysis gracilis, Neomysis integer, Praunus Jlexuosus, P. inermis, Schistomysis ornata, S. spiritus. Pollachius pollachius (L.), pollack: Neomysis integer, Praunus JEexuosus, Schistomysis spiritus, S. ornata. Pollachius virens (L.), saithe : Praunus inermis, Schistomysis ornata, S . spiritus, other species. Trisopterus minutus minutus (L.), poor cod : Erythrops elegans, Praunus Jlexuosus, Schistomysis ornata. Trisopterw esmarki (Nilsson), Norway pout : Erythrops sp., Gastrosaccus spinifer. Trisopterus luscus (L.),bib : Gastrosaccus spinifer, Neomysis integer, Praunus jlexuosus, P. inermis, Schistomysis ornata. Molva molva (L.), ling : Erythrops erythrophthalma. Rhinonemus cimbrius (L.), four bearded rockling : Erythrops erythrophthalma. Gaidropsams mediterraneus (L.), shore ,rockling: Erythrops erythrophthalma. Dicentrarchus labrax (L.), sea, bass : Neomysis integer, Praunus Jlexuosus, Schistornysis ornata.

12.

PREDATORS AND PARASITES

241

Trachurus trachurus (L.), horse mackerel : Mesopodopsis slabberi, Neomysis integer. Boops boops (L.), bogue : Mesopodopsis slabberi. Trachinus vipera Cuvier, lesser weaver : Praunus inermis, Schistomysis spiritus, other species. Swmber (Scomber) scombrus L., mackerel ; Mesopodopsis slabberi, Mysis sp. Gobius niger L., black goby: Praunus jlexuosus. Pomatoschistus microps (Kreryer), common goby : mysids. Pomatoschistus minutus (Pallas), common goby : Neomysis integer, Praunus jlexuosus, Schietomysis ornata. Callionymus lyra L., dragonet : mysids, Neomysis integer. Pholius gunnel1,s (L.),butterfish : mysids. Trigla lucerna L., yellow gurnard 1 Gastrosaccus spinifer, Schistomysis ornata, 8. spiritus. Eutrigla gurnardus (L.), grey gurnard : Erythrops elegans, Neomysis integer, Schistomysis ornata, S. spiritus. Myoxocephalus scorpius scorpius (L.), fatherlasher : Mysis mixta, Praunus jlexuosus. Agonus cataphractus (L.), hook-nose ; Praunus jlexuosus, Xchistomysis ornata. Scophthalmus rhombus (L.), brill ; Neomysis integer. Lepidorhombus whifiagonis (Walbaum), megrim : Erythrops serrata. Psetta maxima (L.), turbot : Gastrosaccus sanctus, Mysis mixta, Paramysis arenosa, Schistomysis spiritus. Arnoglossus laterna (Walbaum), scaldfish : mysids. Pleuronectes platessa L., plaice : Gastrosaccus spinifer, Mesopodopsis slabberi, Neomysis integer, Praunus jlexuosus, P. inermis, Schistomysis ornata, S. spiritus, other species. Glyptocephalus cynogbssus (L.), witch : Mysis mixta, Mysis species. Hippoglossoides platessoides (Fabr.), long rough dab : Erythrops erythrophthalma, Leptomysis lingvura, L. gracilis, Schistomysis ornata, other species. Platichthys jlesus jlesus (L.),flounder : Mesopodopsis slabberi, Mysis mixta, Neomysis integer, Praunus jlexuosus, Schistomysis ornata. Limanda limanda (L.), dab ; Neomysis integer, Praunus JIexuosus, Schistomysis ornata. Solea Eascaris (Risso), sand sole : mysids. The mysids listed above are the commonest species in the northeastern Atlantic with the possible exceptions of Erythrops elegans, which is sometimes locally abundant and Heteromysis formosa, which

242

THE BIOLOGY OF MYSIDS

has never been caught in large numbers. Gastrosaccus spinifer is widely distributed, but may be ineffectively sampled because conventional pelagic or hyperbenthic sledges sre usually employed to sample this species which is in close association with the sediments. It is preyed upon by the epibenthic rays and also by other fish, presumably when it has moved out of and off the bottom and is temporarily pelagic. ,The extent of the predation of any one of these fish species on any one of the mysid species is unknown. I n the Baltic Sea, adult cod feed to a small extent on Gastrosaccus spinifer and Mysis mixta while the flounder consumes some Gastrosaccus spinifer according to Arntz (1978a, b). The early work of Jespersen (1928, 1936)showed that herring around Bornholm fed on Mesopodopsis slabberi, Mysis mixta and Praunus inermis. Mysids were more important however, in the diet of populations of herring near the mouth of the Baltic, around the Danish coasts ; the species eaten were : Gastrosaccus spinifer, Leptomysis gracilis, Mesopodopsis stabberi, Mysis mixta, Neomysis integer, Praunus Jlexuosus, P. inermis, P . neglectus, Schistomysis ornata and S . spiritus. Information on the predatory fish that eat mysids in the northwestern Atlantic is scarce. Brunel (1963b) and Jolicoeur and Brunel ( 1 966) found that cod in the Gulf of St Lawrence feed to some extent on Mysis mixta, while F'itt (1973) states that American plaice, Hippoglossoides platessoides (Fabr.), feed occasionally on mysids on the Grand Banks of Newfoundland. Tattersall (1951)records Mysis stenolepis from the stomach of smelt, Osmerus mordax (Mitchill) off New Brunswick. The white hake, Urophycis tenuis (Mitchill), the longhorn sculpin, Myoxocephalus octodecemspinosus (Mitchill), and the American plaice eat mysids in Passamaquoddy Bay (probably H y s i s stenolepis) (Tyler, 1972). Much more information is available on the predation of mysids whose centres of distribution are south of Cape Cod and is detailed below.

Atlantic Ocean between 40"N and 40"5 The fish that prey upon mysids of the western part of this region will be discussed first. One of the most widespread and common mysids of the eastern coastline of the United States is Neomysis americana and much is known about its predators. It is consumed by shad, mackerel, herring and flounder in Long Island Sound (Whiteley, 1948) and by the common anchovy and sea trout in Delaware (Hopkins, 1965). The following list of predators of N . americana in different coastal regions was obtained from Grant (1962), Richards, (1963), McHugh (1967),

12.

PREDATORS AND PARASITES

243

Markel and Grant (1970), Schaefer (1970), June and Carlson (1971), Sikora et al. (1972), Williams et al. (1974), Stickney et al. (1975), McEachran et al. (1976), Grebe (1978).

Raja erinacea Mitchell, little skate: Cape Cod to Cape Hatteras. Raja ocellata Mitchill, winter skate: Cape Cod t o Cape Hatteras. Raja radiata Donovan, thorny skate: Cape Cod to Cape Hatteras. Raja senta Garman, smooth skate : Cape Cod to Cape Hatteras. Alosa pseudoharengus (Wilson), alewife : New York region. Brevoortia tyrannus (Latrove), Atlantic menhaden : New York region. Merluccius bilinearis (Mitchill), silver hake : New York region. Microgadus tomcod (Walbaum), Atlantic tomcod : New York region. Urophycis Jloridunus (Bean and Dresel), southern hake : Georgia estuaries. Urophycis regius (Walbaum), spotted hake : Georgia estuaries. Menidia menidia (L.), Atlantic silverside : New York region. Morone saxatalis (Walbaum), striped bass : Virginia estuaries. Centropristes striata (L.), black sea bass : Long Island Sound. Pomatomus saltatrix (L.), bluefish : Delaware estuaries. Stenotomus chrysops (L.), scup : New York region. Bairdiella chrysura (Lacepede), silver perch : south-eastern States. Leiostomus xanthurus Lacepede, spot : south-eastern States. Micropogon undulatus (L.), Atlantic croaker : south-eastern States. Stellifer lanceolatus (Holbrook), star drum : south-eastern States. Ammodytes americanus DeKay, American sand lance : New York region. Prionotus carolinus (L.),northern searobin ; New York region. Paralichthys dentatus (L.), summer flounder : Georgia. Paralichthys lethostigma Jordan and Gilbert, southern flounder : Georgia. Scophthalmus aquosus (Mitchill), windowpane : New York region. Pseudopleuronectes americanus (Walbaum), winter flounder : New York region. Neomysis americana is a very important part of the diet of the weakfish according to Stickney et al. (1975) and is also a major portion of the diets of the silver perch, Atlantic croaker and star drum in the estuaries of the south-eastern United States. The striped bass and the bluefish are also major estuarine predators of this mysid (Grant, 1962; Markle and Grant, 1970; Schaefer, 1970). The relative importance of the other fish as predators of this mysid is not known.

244

THE BIOLOGY OF MYSIDS

Much less information is available concerning the predators of other species of mysids occurring along these coastlines. Erythrops erythrophthalma is eaten by the thorny and smooth skates (McEachran et al., 1976). Heteromysis formosa is consumed by the pollack, Pollachius wirens and the northern sewobin and, along with J f y s E ~stenolepis, is also eaten by the scup and black sea bass in Long'Island Sound (Richards, 1963). Tattersall (1951) notes the occurrence of Metamysidopsis munda in the stomach of a gulf kingfish, Menticirrhus litoralis (Holbrook) off Beaufort, North Carolina. The yellow perch Perca LfZawescens (Mitchill), feeds on mysids in the estuaries of Maryland (Muncy, 1962), and the searobins of the family Triglidae also eat mysids but the species consumed were not identified (Ross, 1977, 1978). The predators of Mysidopsis almyra in the brackish lake Pontchartrain, Louisiana are described by Darnel1 (1958). The following list states whether M . almyra was an important or minor constituent of the diet of the fish concerned:

Anchoa mitchilli diaphana Hildebrand, southern bay anchovy : important. Ictalurus furcatus (Lesueur), blue catfish : important. Arius felis (L.), sea catfish: minor. Menidia beryllina (Cope), tidewater silverside : minor. Morone interrupta Gill, yellow bass; minor. Lagodon rhomboides (L.),pin fish: minor. Bairdiella chrysura (Lacepede), silver perch : important. Cynoscion arenarius Ginsburg, sand seatrout : important. Cynoscion nebulosus (Cuvier), spotted seatrout : important. Leiostomus xanthurus Lacepede, spot : minor. Micropogon undulatus (L.), Atlantic croaker : minor. Paralichthys lethostigma Jordan and Gilbert, southern flounder : minor. The Atlantic croaker, Micropogon undulatus, feeds to a notable extent on Mysidopsis almyra in Mississippi Sound and adjacent regions of the Gulf of Mexico (Overstreet and Heard, 1978). No further information appears to be available on the predators of the mysid fauna of the Gulf of Mexico, Caribbean and the coasts of South America. Similarly, little information appears Go be available about the fish preying upon the mysids of the eastern Atlantic between 40"N and 40"s. Longhurst (1957) and Bainbridge (1960) state that a, major predator of Rhopalophthalmus africana and R. longicauda in the Sierra Leone River Estuary is the sciaenid, the gwangura Pseudotolithus elongatus (Bowditch).Further south, off South Africa, Christensen (1978)

12.

PREDATORS AND PARASITES

245

found that the sparids Diplodus cervinus (Lowe) and D. sargus (L.) feed to a small extent on species of mysids, including Mysidopsis similis. Day 1967) records Johnius hololepidotus, the kob, feeding on mysids in the Knysna Estuary in South Africa.

The Mediterranean The following list of predatory fish and the mysids consumed by them was obtained from Reys (1960), Hoenigman (1963) and Haidar (1963).

Raja (Raja)clavata L., thornback ray : Anchialina agilis, Erythrops erythrophthalma, Erythrops sp., Gastrosaccus normani, Lophogaster typicus, Mysideis parva, Mysidopsis didelphys, Pseudomma chattoni, Siriella norvegica, other species. Raja (Dipturus) oxyrinchus L., long-nosed skate : Lophogaster typicus, other species. Raja (Leucoraja)circularis, sandy ray : Lophogaster typicus. Herluccius merluccius (L.), hake : Lophogaster tyiicus. Mullus barbatus L., striped mullet : Anchialina agilis. Trigla lyra L., piper Lophogaster typicus, other species. Trigla lucerna L., yellow gurnard : Gastrosaccus normani, Lophogaster typicus. Aspitrigla cuculw (L.), red gurnard : Anchialina agilis, Erythrops erythrophthalma, Gastrosaccus normani, Leptomysis gracilis, Lophogaster typicus, Mysideis parva, Mysidopsis didelphys, Parapseudomma calloplura, Pseudomma sp., other species. Aspitrigla obscura (L.), shining gurnard : Anchialina agilis,Erythrops erythrophthalma, Erythrops sp., Gastrosaccus normani. Eutrigla gurnardus (L.), grey gurnard : A n c h i a h a agilis, Erythrops sp., Gastrosaccus normani, Leptomysis gracilis, Lophogaster typicus, Mysidopsis didelphys, other species. Trigloporus lastoviza (Brunnich), rock gurnard : Anchialina agilis, Gastrosaccus normani, Lophogaster typicus, other species. Lepidorhombus boscii (Risso), four-spotted scaldfish : Anchialina agilis, Erythrops erythrophthalma, Erythrops sp., Leptomysis gracilis, Lophogaster typicus, Siriella norvegica, other species. Lophogaster typicus was common in the stomachs of several of the fish, but especially those of the thornback ray. It also occurred, along with Boreomysis megalops and other species, in the stomachs of Micromesistius poutassou (Risso), Gadiculus argenteus argenteus Guichenot,

246

TEE BIOLOGY OF MYSIDS

Phycis blennoides (Briinnich), Antonogadus megalokynodon (Kolombatovic) and Symphurus nigrescens Rafinesque in the western Mediterranean (Macpherson, 1978a-c). It is not common in environmental samples because it may live closely associated with the sediment and so i t is not caught by conventional samplers in any great numbers. It must, however, be a relatively numerous species at least in some localized areas. The Black and Caspian Seas Sturgeon feed to some degree on Paramysis species in the Sea of Azov (Savchuk, 1975) while young beluga, Huso huso (L.), feed on Paramysis baeri and P. intermedia in the northern Caspian Sea (Belyayeva et al., 1972). Mikhman and Romanovich (1977) state that the Azov anchovy, Engraulis encrasicolus (L.), feeds on mysids but they did not identify them. Young Percarina demidofli Nordmann consume mysids in the Gulf of Taganrog (Mikhman, 1978). Mysids assume relatively greater importance in the diets of fish in the Black and Caspian Seas than in other regions because there are marked decreases in the variety and numbers of decapod shrimps present in these areas of decreased salinity. Bacescu (1938) describes the extent of this predation on the various endemic mysids in considerable detail.

North Pacific, north of about 40"N

No information has been found regarding fish that prey upon mysids of this region, except in the coastal areas of the north-western United States and in the western Pacific. Neomysis species are eaten by the Pacific sanddab, Citharichthys sordidus (Girard), and the petrale sole, Eopsetta jordani (Lockington), off the coast of Oregon (Kravitz et at., 1976). Neomysis kadiakensis off the coasts of Washington State is eaten by the flathead sole, Hippoglossoides elassodon Jordan and Gilbert, the stony flounder,, Platichthysl stellatus (Pallas), and the sand sole, Psettichthys melanostictus Girard (Miller, 1967, 1970). A further species, Archaeomysis grebnitzkii, is reported as a major food of the fish of the Columbia River by Haertel and Osterberg (1967). An ecological study of Lake Washington showed that Neomysis mercedis (wrongly identified as N . awatschensis; see Holmquist, 1973) is of varying importance in the diets of the prickly sculpin Cottus asper Richardson, three spined stickleback Gasterosteus aculeatus L. longfin smelt Spirinchus thaleichthys (Ayres) and the yellow perch Perca fivescens (Mitchill) (Eggers et al., ~

12. PREDATORS

AND PARASITES

247

1978). This population of Neomysis mercedis has declined over the period 1962-1964 to 1975: Eggers et al. suggest that production of benthos decreased owing to a diversion of sewage wastes and the fish subsequently preyed more heavily upon the mysids causing a reduction in their numbers. In the western part of this region, Skalkin (1959) found that flounders in the Sea of Okhotsk included mysids in their diet. Further south, on the coasts of Japan, Enomoto (1964)states that Parastilomysis paradoxu is eaten by Cocius crocodilus (Tilesius) and Bembras japonicus Cuvier and Valenciennes. The coastal Neomysis intermedia is consumed by smelts (Murano, 1963a),while N . czerniavskii is occasionally eaten by juvenile chum salmon in regions of Japan (Okada and Taniguchi, 1971).

Pacijc Ocean, between about 40"N and 40"5 Neomysis mercedis, incorrectly identified as N . awatschensis by earlier authors (Holmquist, 1973; Simmons et al., 1974), is a dominant species in the Sacramento-San Joaquin Delta, California. The predators of this species have been studied by Heubach et al. (1963), Ganssle (1966), Radtke (1966), Sasaki (1966a, b), Stevens (1966a, b), Turner (1966a, b) and Thomas (1967). They are listed below with an indication of whether the mysid is an important or minor part of their diets. Acipenser medirostris Ayres, green sturgeon : important. Acipenser transmontanus Richardson, white sturgeon : important. Alosa sapidissima (Wilson), American shad : very important. Oncorhynchus tshawytscha (Walbaum), chinook salmon : important. Ictalurus catus (L.), white catfish : important. Ictalurus melm (Rafinesque),black bullhead : minor. Ictalurus nebulosus (Lesueur), brown bullhead : minor. Ictalurus punctatus (Rafinesque), channel catfish : minor. Morone saxatilis (Walbaum), striped bass : important. Archoplites interruptus (Girard), Sacramento perch : unknown. Lepomis cyanellus Rahesque, green sunfish : unknown. Lepomis gulosus (Cuvier), warmouth : important. Lepomis macrochirus Rafinesque, bluegill : minor. Micropterus salmoides (Lacepede), largemouth bass : minor. Pomoxis annularis Rafinesque, white crappie : unknown. Pomoxis nigromculatus (Lesueur),black crappie : very important. Platichthys stellatus (Pallas), starry flounder : minor. The fact that Neomysis mercedis is important in the diet of many of 1 I: , @@oL ~ ~ ) O O ~ O U o]]o--- -4.-- 4 C IJ 3 . species only allows estimates of the predation pressure upon it to OD]XX

I

: I

I

248

THE BIOLOGY OF MYSIDS

be made if considerable information is available about the biomasses of the mysids on the one hand and of the fish on the other. This information does not appear to be available but it is very probable that these populations of N . mercedis are subject to considerable predation pressure, a t least seasonally. The Pacific sardine feeds t o a minor extent on mysids but the species have not been identified (Hand and Berner, 1960). An ecological group of fish associated with the kelp beds of the Californian coasts prey to a considerable degree on the swarms of mysids living in this habitat (Hobson and Chess, 1976; Love and Ebeling, 1978). Two such mysids are Acanthomysis sculpta and Siriella pacifica which are eaten by fish such as the kelp bass, Paralabrax clathratus (Girard), the salema Xenistius califormiensis (Steindachner), the queenfish, Scriphus politus Ayres, the kelp perch, Brachyistius frenatus Gill, the walleye surfperch, Hyperprosopon argenteum Gibbon, the kelp rockfish, Sebastes atrovirens (Jordan and Gilbert), the blue rockfish, S. mystinus (Jordan and Gilbert) and %heolive rockfish, S. serranoides (Eigenmann and Eigenmann). Coral reefs form a similar type of habitat where mysids and fish live in close proximity. The formation of swarms by the mysids, as discussed previously, appears t o confer protection on them from predation during the day. On the other hand, many of the reef fish aggregate for their own protection from diurnal predators. Hobson and Chess (1978) found that the heaviest predation on mysids was effected by nocturnally feeding fish. The following species of fish of the coral reefs of Hawaii prey t o some extent on mysids (Hobson, 1974): Brotula multibarbata Temminck and Schlegel, Pranesus insularum (Jordan and Evermann), Holocentrus diadema Lacepede, H . xantherythrus Jordan and Evermann, Myripristis amaenus (Castelnau), M . kuntee Cuvier, M . murdjan (ForskAI), Priacunthus cruentatus (Lacepede), Apogon menesemus Jenkins, A. snyderi Jordan and Evermann, Hemitaurichthys thompsoni Fowler, Chuetodon corallicola Snyder, C. milearis Quog and Gaimard, Chromis ovalis (Steindachner) and C. verater Jordan and Metz. Two studies of the diets of the fish of the coral reefs of the Marshall Islands have been made (Hiatt and Strasburg, 1960; Hobson and Chess, 1978). The following species feed to some extent on mysids: Hyporhumphus laticeps (Gunther), Myripristis pralinius Cuvier, M . violaceus, Apogon savayensis Gunther, A. novae-guineae Valenciennes, Dascyllus reticulatus (Richardson), Chromis agilis, C. atripectoralis Welander and Schultz, C. caeruleus (Cuvier and Valenciennes), Abudefduf leucopomus (Lesson), Pomacentrus pavo (Bloch), Labroides dimidiatus (Cuvier and Valenciennes). Immature copper sweeper, Pempheris schomburgki

12.

PREDATORS A N D PARASITES

249

Muller and Troschel, eat mysids among the coral reefs of the Virgin Islands but Gladfelter (1979) did not identify the species consumed. In the southern Pacific, off New South Wales, Australia, the scorpaenid Centropogon australis (White) feeds t o a marked extent on Australomysis species (Bell et al., 1978). There appears to be no further information available on the fish that predate on the mysids of the Pacific Ocean.

Indian Ocean Little information can be located regarding the fish that are predatory upon the mysids of this ocean. Whitfield and Blaber (1978), examining the feeding of fish in Lake St Lucja in south-west Africa (28"S), found that Terapon jarbuu (ForskAl) and Johnius belengerii (Cuvier) feed on Mesopodopsis africana. Birds may contribute significantly to the predation of mysids in certain regions. Surface swarms of epipelagic species will be accessible to them and of course most swarming littoral species will be subjected to a t least occasional predation by birds. Tattersall (1951) notes that Antarctomysis maxima in the Antarctic is recorded from the stomach of the penguin Pygoscelis papua (Forster) while Archaeomysis grebnitzkii has been recorded in the Aleutians in the stomach of an eider duck. Tattersall and Tattersall (1951) have summarized $he few early records of birds feeding on mysids. Wading birds in Norfolk, England have been observed feeding on Neomysis integer and Praunus flexuosus. Mysis oculata was reported as important in the diets of kittiwakes Rissa tridactyla tridactyla L., fulmars Fulmaris glacialis glacialis L., Arctic terns Sterna paradisea Brunn, northern eiders Somateria mollissima borealis Brehm., Mandt's guillemot Uria grylle mandtii Mandt, little auks Plotus alle L. and Spitzbergen puffins Fratercula arctica naumanni Norton in western Spitzbergen by Hartley and Fisher (1936). More recently, Zelikman and Golovkin (1972) found that the little auk, Plotus alle L., included Mysis oculata in its diet in Novaya Zemlya. Mysids are eaten by the crested auklet, Aethia cristatella (Pallas), the least auklet, A . pusilla (Pallas),and the parakeet auklet, Cyclorrhynchus psittacula (Pallas)in Alaska (Bedard, 1969). The sand-burrowing species of mysids of the genus Gastrosaccus are accessible to birds a t low tide. Moran and Fishelson (1971) found that G. sanctus on Israeli beaches was preyed upon by the plovers Charadrius alexandrinus L. and C. hiaticuEa L. Tattersall and Tattersall (1951) cite Dana's record of the occurrence of the bat,hypelagic Eucopia australis in the stomach of a

250

THE BIOLOGY OF MYSIDS

penguin when it ate a (vertically migrating) fish. Holthuis and Sivertsen ( 19671, examining the stomach contents of the albatross Diomedea chlororh,ynchus Gmelin taken near Tristan da Cunha, found that they contained the bathypelagic Chalaraspidum alatum, Gnathophausia ingens and G. zoea; they suggest that deep water must presumably upwell somewhere in this region to make these species accessible t o the birds. Nemoto (1959) found a young G. gigas in the stomach of a fin whale in the North Pacific and it may be that species in this genus sometimes migrate to the surface layers. Several invertebrates prey on mysids. Lebour (1923) suggests that the ctenophore Pleurobrachia pileus (Fabr.) eats mysids. The sanddwelling Gastrosaccus pammodytes in South Africa is ,apparently eaten on occasions by the gastropod snails of the genus Bulla, especially if the snails are starving (Brown, 1964).The other recorded invertebrate predators are all crustaceans. The bathypelagic ostracod Gigantocypris muelleri Skogsberg, according to Moguilevsky and Gooday (1977), feeds to a small extent on Eucopia unguiculata and E. grimaldii. Matsudaira et al. (1952) describe isopods as a natural predator of Gastrosaccus vulgaris. Lagardbre (1972, 1977a, b) and Inyang (1978) found that the following decapods included mysids in their diets bu% the species were not identified : Pasiphaea tarda Krpryer, Palaemon adspersus var. fabricii (Rathke), Plesionika heterocarpus (Costa), Philocheras echinulatus (M. Sars), Parapagurus pilosimanus Smith. Lagardbre ( 1973, 1977b) found that Dichelopandalus bonnieri Caullery included mysids as a minor item in their diet in the Gulf of Gascogne; he recognized portions of Lophogaster typicus among the stomach contents but other species were probably also present. Crangon crangon (L.), in the littoral environment, is known to consume Mysis species and Mesopodopsis slabberi according to Bacescu (1938) and Tattersall and Tattersall (1951). The related species Crangon septemspinosa (Say) includes Neomysis americana in its diet in Delaware Bay (Price, 1962; Richards and Riley, 1967). LagardBre (1972) found that Parapenaeus longirostris (Lucas)included the following mysids in its diet : Anchialina agilis,Gastrosaccus sp., Heteromysis formosa, Leptomysis gracilis, Lophogaster typicus and Mysideis sp. He further found Paramblyops rostrata in the stomachs of Aristeomorpha foliacea (Risso) and Solenocera membranacea (Risso) and, along with Boreomysis species, in the stomachs of Plesopenaeus edwardsianus (Johnson). Lagardbre (1977b) noted Gastrosaccus lobatus among the stomach contents of Plesionika heterocarpus, Solenocera membranacea, Philocheras echinulatus (M. Sars), Nephrops norvegicus (L.), Munida sarsi ,and Polycheles typhzops Heller. In addition, P. typhJops ate Gnathophausia ,zoea, Boreomysis arctica and '

12.

PREDATORS AND PARASITES

251

Parapseudomma calloplura; Munida perarmata Milne-Edwards and Bouvier ate Boreomysis species and Geryon longipes Milne-Edwards ate Cnathophausia zoea. Clearly, mysids are subjected t o predation by a great variety of organisms. This is hardly surprising because many species occur in vast numbers in restricted sea areas. These areas have endemic fish populations, others are nursery grounds for young fish. The body size of most coastal species of mysids is in the same range as that of food organisms of very many fish. The mysids, however, have developed various means of protection against predators. Many are cryptically coloured and blend with their environment. Others form swarms and so produce scattered concentrations of individuals that may reduce the chance of detection by fish (Clutter, 1969). Yet others have developed peculiar behavioural traits : an example is Mysidium gracile, schools of which, when threatened by a predatory fish, crowd into the nesting caves of several species of pomacentrid fish (Emery, 1968). It is very difficult to assess the impact of predation upon a population of mysids. The populations of many species show large changes in apparent biomass over relatively short periods of time. Increases in the apparent size of a population can often be accounted for by aggregation of a previously disaggregated population or through natural increases in population size by production of young. Decreases in population numbers are more difficult to account for satisfactorily. An aggregated population can become disaggregated and inhabit a larger portion of the environment. This situation is usually difficult to quantify and so obtain confirmation that no abnormal morbality through predation has occurred. There are decreases in numbers within populations through natural mortality, either age-related or at mating, in the case of males, or at egg laying, in the case of females. Mortality associated with mating or egg laying may not in fact be directly caused by these events, e.g. death through exhaustion. The mysids may be more readily caught by predators as a consequence of these events. Much more information is required 60 assess the roles of predators in the population dynamics of mysids. A wide range of parasites has been reported in mysids, the most frequently described' being the ellobiopsid species of the genus Thalassomyces. These have been classed as Protista (incertae sedis) by Wing (1975) although Galt and Whisler (1970) describe the production of flagellated spores by Thalassomyces marsupii and suggest affinities with the Dinoflagellata. The parasite (Fig. 63) normally protrudes from the carapace but can alternatively occur within the marsupium or on the abdominal segments. It consists of a sub-integumental body that

t S

C

E

D

FIG.63. Parasites of mysids. A, the trophomeres of a Thah8omyce.5 sp. protruding from the carapace of mysid; B,enlarged drawing of the trophomeres showing ( g ) ,the gonomeres and (s),spores emerging from a terminal gonomere. C, a femalechoniostomatid copepod with egg sacs. D, an epicaridean isopod of the genus A8pidophryw attached t o the carapace of a mysid and enlarged in dorsal view. E, juvenile epicaridean isopods of the genus Dajus in dorsal and lateral view. (C, after Mauchline, 1968; D and E, after Sars, 1898).

12.

PREDATORS AND PARASITES

253

probably develops.from a “spore” ingested by the host (Kane, 1964; Hoffman and Yancey, 1966; Mauchline, 1966). The parasite develops and grows outwards through the integument, its reproductive organs, the trophomeres, being the external evidence of its presence within the mysid. The trophomeres are segmented, consisting of a chain of gonomeres within which the spores develop (Fig. 63). The identification of the species of ellobiopsid present in a mysid, or any other crustacean, is difficult, the characters used being the number of trophomeres, the numbers of gonomeres per trophomere and the shape and proportions of these organs (Wing, 1975). These characters are all variable. Multiple infections are known but rare and only one group of trophomeres is usually present. No detailed study of their occurrence has been made in any mysid and much remains unknown about their physiology, reproduction and life history. Their occurrence in euphausuds and decapods has been examined more frequently (Mauchline, 1980). The internal absorptive or haustorial system can penetrate the host’s nervous system, prevent or hinder development of the host’s reproductive system or cause other abnormalities. Nothing of these aspects is known in mysids. Various species within the genus Thalassomyces have been described (Vader, 1973) and they infect species of Amphipoda, Mysidacea, Euphausiacea and Decapoda. The various species known to infect mysids are listed in Table XXXIX. As can be seen, they infecti mysids from a number of genera but why they occur in one species of a genus and not in another is unknown. The lest species of ellobiopsid listed in Table XXXIX, Ellobiocystis mysidarum, is at present included in the Ellobiopsidae although its affinities with these organisms is in question. It has no sub-integumentary root and is a non-parasitic epibiont resembling a single trophomere of a Thulassomyces species. It is smaller in size and attached singly or in clusters, usually to the mouthparts of the host. Various other protozoans are known to occur in or on mysids. Species of suctorians identified as belonging to the general OphryodendTon Claparhde and L. and Paracineta Collin were identified on Siriella jaltensis gracilipes by Hoenigman ( 1960). Tattersall and Tattersall (1951) refer to earlier records of Acineta tuberosa (Miiller) infecting Mysis relicta, Epistylus species on Praunus neglectus and Zoothamnium species on Praunus Jlexuosus and Neomysis integer. This latter species is also frequently infected with apostome ciliates. Internal protozoan parasites of mysids have onlyheen examined by Landau et al. (1975) who found sporozoites of a coccidian parasite of the fish Gymnothorax moringa (Cuvier); the species of mysid was not named.

TULE XXXIX. ELLOBIOPSID PARASITES OF MYSIDS IN DIFFERENT REGIONS Ellobiopsid Thalassomyces boschmai (Nouvel)

Thalassomycea fasciatzls (Fage)

Thalmaomyces bitowskii (Hoenigman) T h a h s m y c e a nouveli (Hoenigman) Thalassomycea albatrossi Wing Thalassomyces sp. Ellobiocystis rnysidarum Coutihre

Mysid Acanthmysis pseudomacropsis Gastrosaccus lobatus L e p t m y s i s gracilis Meterythrops robuata N e m y s i s kadiakensis Cnathophausia zoeu clulthophausia gigas Cnathophamia gracilis Cnathophazcaia ingens L e p t m y & t megalops Anchialina agilis Stilomysis major Siriella clausii Antarctomysis maxima

Region

References

N.E. Pacific Mediterranean Mediterranean N.E. Pacific N.E. Pacific Atlantic, Pacific E. Atlantic Pacific Atlantic, Pacific Adriatic Mediterranean Korea Mediterranean Antarctic

11,12 8 5, 6, 8 12 11,12 1, 2 2 10 1, 2, 10 7 4, 8 12 9 3

References:, Fsge, 1936, 1941; 2, Nouvel, 1941; 3, Boschma, 1949; 4, Hoenigmen, 1954; 5, Nouvel, 1954; 6, Nouvel and Hoenigman, 1955; 7, Hoenigman, 1960; 8, Hoenigman,1963; 9, Hoenigman, 1966; 10, Pequegnet, 1965; 11, Wing, 1935; 12, Wing, 1975.

12.

255

PREDATORS AND PARASITES

The occurrence of parasitic worms in myaids has been rarely observed. Brownell ( 1970) concluded, on experimental evidence, that Mysis relicta is not a vector of the acanthocephalan Echinorhynchus salmonis Muller, a parasite of the lake trout. Amin (1978) found one Mysis relicta in Lake Michigan infected with a procercoid of the cestode worm Cyathocephalus truncatus Pallas, 1781. Schistomysis spiritus was shown by Shotter (1970, 1972) to be a second intermediate host of the digenean trematode, Podocotyle atomon (Rudolphi) known as the whiting worm. Tattersall and Tattersall (1951) refer to Acanthomysis strauchi as the intermediate host of the cestode Amphilina foliacea (Rudolphi), a parasite of the sturgeon. Bacescu (1954) notes the presence of cysts in the musculature of Paramysis kessleri sarsi. TABLEXL. CHONIOSTOMATID COPEPOD PARASITES OF MYSIDS (Tattersall and Tattersall, 1951 ; Mauchline, 1969b) Gopepod

Mysidion abyssorurn Mysidion commune Hansen Aspidoecia norrnani Giard and Bonnier

My& Erythrops abyssorum Erythrops abyssorurn Erythrops serratu Parerythrops obesa Erythrops abysaorurn Erythrops elegans Erythrops erythrophthalma Erythrops microps Erythrops serrata

Location

marsupium marsupiurn marsupium marsupium abdominal segments eyes, marsupium

There are several parasitic crusbaceans that infesb mysids. Three species of choniostomatid copepods (Table XL), have been recorded. The two Mysidion species occur in the marsupia of Erythrqs and Parerythrops species, where they are attached to the ventral integument. They are roughly spherical in shape (Fig. 63), the female having ovisacs protruding from her body. The general body form of Aspidoecia norrnani is similar but it differs from that of Mysidion species in having no maxillipeds. Further, although it does occur attached to the ventral thoracic region of the mysids, it is more commonly found attached to the abdomen and even the carapace and eyes. Species belonging to two families of epicaridean isopods parasitize mysids (Table XLI). The females are often relatively large in body size (Fig. 63) although juvenile stages and the smaller sized males also occur within the marsupium where they live freely among the larvae of the mysid. Many of these parasites, if attached to the dorsal regions of the

TABLE XLI. EPICARIDEAN ISOPOD PARsSITES

Iaopod Asconiscidae Asconiacw simplex G. 0. Sars Dajiidae Arthrophryxw beringanua Richardson Aspidophryxua frontalis Bonnier Aspidophryxua peltatus G. 0. Sars

Uajus mysidis Kr0yer Dajua siriella G. 0. Sars

Holophryxua a h c e n s i s Richardson Notophryxus clypeatua G. 0. Sars

Mysid

OF

MYSIDS

Location

Boreomysis arctica

References

marsupium

3

Eucopia australis

3

Siriella norvegica

rostrum

3, 6

Erythrops elegana

carapace

3

Erythropa erythrophthalma Erythrops micropa Erythropa serrata Mysidopsia didelphys Parerythrops obeaa Mysia mixta Mysis oculata Siriella thompaonii Siriella aequiremis Siriella inornata Siriella vulgam'a My& polaria

carapace

3

carapace carapace carapace carapace marsupium marsupium marsupium marsupium

3 3, Q 3, 11 3 3 3 3 2 2 10

Parapseudomma calloplura Paeudomma aflne Pseudomma roaeum Amblyops abbreviatcc

marsupium

3

Notophryxus ovoides G. 0. Sars Prodajus gaatrosacci Pillai Gaatroaaccua simulans Prodajus lobiancoi Bonnier GaatToaaccua Eobatua Gaatroaaccua n o m a n i Prodajua ostendensis Ga.strosaccua apinifer Gilson Prodajua ovatua Pillai Gaatroaaccua muticua Gaatroeaccua eimulana Streptodajua equilibrans Boreomysis aibogae ~

-

2

carapace carapace abdomen

12 3 3

marsupium

8 4, 6, 7 3, 5 1, 3

-

marsupium

8 13 2 ~~~

~

~

References: 1, Gilson, 1909; 2, Tattersall, 1951; 3, Tattersall and Tattersell, 1961; 4,Nouvel, 1951b;6,Teberly, 1964a,b;6, Hoenigman, 1958b, 1963;7,Hoenipan, 1960; 8, Pillai, 1963c; 9, Mauchline, 1968; 10, Geiger, 1969; 11, Mauchline, 1970b; 12, Mauchline, 1971g; 13, Balasubrahmanyan and Prince, 1976.

12.

PREDATORS AND PARASITES

257

integument, become detached during the processes of catching the mysids. Consequently, they may be more common than present evidence suggests. They may be overlooked among the larvae in the marsupium or, if noticed, considered as intruders that have entered the marsupium when the mysid was in the crowded bucket of the plankton net. Epiphytic diatoms and algae often occur on the integuments of mysids. Hoenigman (1958b, 1960) describes Diamysis bahirensis, Leptomysis lingvura, Siriella clausii and S . jaltensis gracilipes with diatoms of the genus Licmophora attached to them; he identified Licmophora dalmatica Gr. on Diamysis buhirensis. Schistomysis spiritus frequently has Licmophora lyngbyei and Navicula species attached to its eyes and other parts of the integument; Ectocarpus species also frequently occurs on the carapace and eyes of this species (Mauchline, 197lg). Hoenigman (1958b) recorded the hydroid coelenterate, Clythia johnstoni Alder, attached to one of the uropods of Siriella jaltensis gracilipes.

CHAPTER 13

MYSIDS IN THE MARINE ECONOMY The array of known predators of mysids indicates their general importance in the food webs of the seas. The mysids are generally mixed feeders although considerable variation exists between different species and also within a species a t different times and places. Praunus jlexuosus can feed on a diet of algae and dead animal material, be a strict carnivore feeding on cyclopoid or harpacticoid copepods or a general omnivore, while a species such as Mysidopsis gibbosa is more strictly carnivorous. Most coastal species utilize organic detritus (or debris) to a considerable extent. This debris includes terrigenous material entering the sub-littoral from freshwater streams and rivers. Brown (1964), studying the food relationships on an intertidal sandy beach in South Africa, suggests that Gastrosaccus psammodytes consumes detritus and is itself eaten by teleost fish. North-eastern Atlantic species of the genera Praunus, Paramysis, Schistomysis and the species Neomysis integer and Mesopodopsis slabberi also feed on organic debris and are the prey of teleost fish. It is possible, therefore, that littoral mysids mobilize the larger organic particles in the environment. Much of this material arises from two sources. The first is the terrigenous environment and the second is through the breakdown of macroalgae. Consequently, these mysids are involved in a short food chain: organic debris-tmysids-tfish The harpacticoid and cyclopoid copepods, feeding on phytoplankton and a smaller size spectrum of organic debris, are important prey of mysids. The biomass of littoral mysids present in a lagoon or over a semi-sheltered sandy beach is often large although irregularly distributed. The irregularity of their distribution is correlated with avoidance of wave action and availability of food, assuming that the total environment has the characteristics of salinity and temperature preferred by the species in question. Consequently, the impact of the mysids on the quantity of organic debris in the area may be quite significant, although no quantitative information on this is available. Mysids, of course, do

13.

MYSIDS IN THE MARINE ECONOMY

259

not live in isolation from other organisms and large numbers of amphipods frequently occur in the same samples. The amphipods attack the mysids within the sampler but there is no quantitative information about predation of mysids by amphipods in the natural environment. Gammarus species bite the mysids, often removing a portion of the abdominal musculature. The amphipods also feed on organic debris and may compete with mysids under certain conditions. The lititoral fish usually have mixed diets of molluscs and crustaceans, including mysids in many instances, so that the mysids are involved in more or less complex food webs in littoral and immediately sub-littoral situations. The biomass of mysids and its spatial distribution within the environment affects bheir availability to predators, especially fish. The biomass present in various sectors of the marine environments varies greatly. Murano et al. (1976) found 0-05-0.15 g wet weight mysids per 1000m3 of water off Central Japan. Maynard et al. (1975) measured biomasses of the order of 0.01-0.10 g wet weight per 1000 ms in the 0-1200 m water column in the region of Hawaii. The biomass of mysids in the California Current varies considerably, not only beeween offshore and coastal regions, but also within these regions at different seasons; Isaacs et al. (1969) recorded up to 2-3 g wet weight mysids per 1000 m3 water on one occasion, in April, 1956. Vinogradov (1970a) found that the biomass of mysids in the water column of the Kurile-Kamchatka Trench ranges from 0.05-3-Og wet weight per 1000mS water, the highest biomass occurring in the 500-1500 m depth layer. The modal depth of occurrence of the biomass of mysids is deeper t,han that of euphausiids in the oceanic water column (Vinogradov, 1970a; Vinogradov and Parin, 1973; Murano et al., 1976). This is illustrated in Mauchline (1980). Euphausiids and mysids appear to be mutually exclusive in their distributions. Mysids occupy the estuarine, littoral and sub-littoral environments where euphausiids are absent. In the neritic environment, mysids are predominantly hyperbenthic while euphausiids are predominantly pelagic. A few oceanic species of mysids are epipelagic but the bulk of the biomass is mesopelagic or bathypelagic. Oceanic euphausiids are predominantly epipelagic although a few are meso- and bathypelagic. Euphausiids and mysids were originally classified together as the Schizopoda because of many basic similarities and would probably be direct competitors if they occupied the same regions of the marine environment. The biomasses of mysids in offshore areas have been measured through the use of pelagic nets. Much greater biomasses per unit volume of water occur in restricted regions of the littoral and neritic environments, especially at the sediment-water interphase and when swarming

260

THE BIOLOGY OF MYSIDS

species are present. The total biomass of these populations, in terms of a fishery resource, is small although many are exploited commercially. Morgan and Beeton (1978) found that populations of the freshwater Mysis relicta, which is frequently described as a benthic organism in Lake Michigan, average approximately 200 individuals per m2 of sediment surface; their samples were collected by an epibenthic sledge. This represents m approximate biomass of 2-20 g/m2,depending on the body size of the individuals. The distance between individuals, assuming they are evenly distributed over the surface of the sediment, is 7 cm, that is approximately equal to the square root of the area occupied by a single individual. The distance between individuals in swaxms is about 2 cm (Table XXX, p. 179) or about 2500 individuals/ma, assuming that they are distributed in one layer over the sediment. Swarms, however, although frequently laminar in form are not composed of a single layer of individuals but of several layers. The density found by Morgan and Beeton in populations of Mysis relicta is similar to those of several neritic marine species in the hyperbenthic habitat. Leptornysis g r a d i s , Erythrops serrata and Mysidopsis didelphys in coastal regions of Scotland occur a t such densities. Such densities are probably quite common among aggregations of neritic hyperbenthic species, including the larger offshore populations, for example those of Georges Bank, of the estuarine and coastal Neomysis americana. Richards and Riley (1963) found thab N . americana represents 13-66% by number and 0-2-1.0% by weight of the total epifauna h regions of Long Island Sound. The mean standing crop ranged from 2.6 t,o 21.6 individuals/m2 or 2-12 mg/m2. High densities will, however, tend to occur irregularly in time and space, being dependent upon such factors as the physical, chemical and topographical characteristics of the environment and the behavioural patterns of the populations, e.g. the occurrence of restricted distributions or breeding aggregations. Littoral and sub-littoral swarming species occur a t biomasses of the order of 20-200 g/m2 depending upon the sizes of the individuals concerned. The swarms, however, with total biomasses in this range, are often small in size, and irregularly distributed within their zone of occurrence. Such swarms are exploited commercially especially in tropical and subtropical regions but the extent of this exploitation is not known. Part of the reason for this is that, as Omori (1978) points out, the species concerned are often unknown simply because they have not been examined by a taxonomist. Omori states that several species of mixed groups of species are fished in local regions of China, Korea and south-eastern Asian countries but no descriptions, quantitative or qualitative, of these fisheries are available. The mysids are used to make

13.

MYSIDS IN THE MARINE ECONOMY

261

shrimp paste and sauces. Some thousands of tons of Neomysis intermedia, N . japonica dnd Acunthomysis mitsukurii are harvested in Japan each year. The most important species is Neomysis intermedia from brackish lakes; it is boiled, dried in the sun and frequently marketed as a preserved cooked food known as tsukudani. Tattersall end Tattersall (1951) review instances of commercial exploitation of mysids. Praunus Jlexuosus was collected in the Channel Islands and made into a paste called cherwe for use as bait for catching mullet. Mesopodopsis orientalis was mixed with smaller numbers of Gangemysis assimilis and sold as Kada chingri on the markets of Calcutta. Mesopodopsis orientalis was eaten elsewhere in India, where it was known as Sridhar. No recent information about these fisheries for mysids is available and so no confirmation of their continuance has been obtained. Mysids have recently been used in fish farming projects as a live food resource, especially in freshwater fisheries. Mysis relicta has been introduced t o various lakes in Sweden (Furst, 1965) and in Canada and the United States (Sparrow et al., 1964; Linn and Frantz, 1965; Hanson, 1966; Schumacher, 1966; Stringer, 1967). The following species have been introduced to various reservoirs and rivers of the PontoCaspian region according to Zhuravel (1959, 1960a, b), Tskhomelidze and Sergeeva (1966), Boroditch and Havlena (1971), Boroditch (1978) and Mordukhai-Boltovskoi (1979) ; Hemimysis anomala, Limnomysis benedini, Paramysis baeri, P . intermedia, P . lacustris and P . ullskyi. Most of these introductions appear to have been successful, in some cases probably too successful. Richards et al. (1975) have described the disruption of the ecosystem of Lake Tahoe, California following the introduction to it of Mysis relicta and kokanee salmon fry and Zyblut (1970) examined similar changes following the introduction of M . relicta to Koohnay Lake, British Columbia. According to Northcote (1972, 1973), M . relicta was introduced to Kootenay Lake in 1949 and had become established and abundant by the early 1960s. He concludes that the mysids are probably responsible for the increased size and growth rates of the kokanee salmon and mountain whitefish. There was no evidence however, that medium sized trout had benefited. Mysis relicta is a potential predator of the native zooplankton populations. Ivanov (1972) discusses the benefits to fish production of stocking ponds or reservoirs with mysids and shows that increased production is obtained. Ogle and Price (1976) suggest that mysids may be a suitable food organism for use in prawn culture ; $hey found that the commercial brown shrimp Penaeus aztecus grew at comparable rates when fed on a diet of Mysidopsis almyra as when fed a diet of Artemia nauplii.

262

THE BIOLOGY OF MYSIDS

Methods of cultivating mysids have not been described in any detail, with the exception of the work bf Murano (1966c) on Neomysis intermedia. He successfully reared it through several generations in the laboratory. Satisfactory diets included larvae of Artemia and freshwater cladocerans but the presence of mud or sandy mud on the bottom of the culture vessels was also found to be necessary. Many species are easily maintained in the laboratory for experimental purposes although no published accounts of continuous culturing have been found. Lawler and Shepard (1978) describe methods of eradicating hydrozoan pests in mysid cultures. Most culturing has been simply carried out by introducing a sample of juveniles and adults to a restricted location such as a pond, lake or river. Ivanov (1972), for example, used seasonal fish ponds. Consequently, this culturing is done in semi-natural environments and does not necessarily involve simultaneous artificial production of food as a continuous culturing system in a laboratory would require. Future work may demonstrate the advantages of artificial culture of mysids as a fish or prawn food, although Blegvad (1922) suggests that the potential production of littoral amphipods makes them more attractive for this purpose. There are two other instances of transplantation of mysids. Zhuravel ( 1 972) introduced the Caspian species Pararnysis kowalewski (now P. lacustris),along with some amphipods, into several areas of Lithuania and Estonia to increase the supply of food for fish. This transplantation was in 1953 and the mysids and amphipods became established in the Kaunas reservoir on the Newman River in Lithuania. This river flows to the Baltic and P. Zacustris first occurred in the Baltic Sea off Estonia in 1955 and has since been found at other locations. The second instance of transplantation is that of Praunus jiexuosus, endemic to bhe northwestern European coastal regions. It was first reported in the Western Atlantic in New England coastal waters by Wigley (1963) but has since extended its range and become common on the coastlines of Massachusetts, New Hampshire and Maine. Pezzack and Corey (1979) record it in Passamaquoddy Bay. There is no record of its introduction and Wigley and Burns (1971) suggest that it may have been transported among fouling organisms on ships. Mysids are excelIent experimental organisms. They are common in littoral regions and do not require expensive ship-time and sampling equipment for their collection. They can be maintained easily in the laboratory (Lawler and Shepard, 1978). Biesinger et al. (1976) used Mysis relicta to test the toxicity of polyelectrolytes while Jacobs and Grant (1974) examined the reaction of Neomysis americana to Kraft mill effluent. The estuarine species, Mysidopsis bahia, was cultured by

13.

MYSIDS IN THE MARINE ECONOMY

263

Nimmo et al. (1978) a t temperatures of 25-28°C. Its generation time was 17 days. They tested its sensitivity to various concentrations of cadmium over its entire generation time. The mysid was sensitive to much lower concentrations of cadmium than those reported for other organisms. The LC 50 was 15.5 pg Cd/l compared with an LC 50 of 320 pg Cd/l for hermit crabs and sand shrimps. The toxicity of the insecticide lindane was examined in M . bahia by Schimmel et aE. (19771, the 96 h lindane LC 50 being 6.3 pg/l. Consequently, some species of mysids may be extremely useful experimental organisms in studies of the potential impact of various pollutants on the environment. Mysis etenolepis was found to be able to utilize cellulose, probably through the presence of a gut microflora (Foulds and Mann, 1978). It assimilated sterile raw cellulose, with efficiencies of 30-50%, efficiencies that are surprisingly high and could be of interest in the context of wood pulp mill effluents. There is also a possibility that a historical record of past densities of mysids may exist in some sea areas. Enbysk and Linger (1966) found that the statoliths of various species persisted in the sediments of continental shelves. They are apparently absent in samples of slope sediments but this may reflect the lower densities of mysids occurring in the overlying water column. Their abundance in coastal sediments ranged from 1-5 statoliths/g dry weight of sediment. An addendum to this review will be found starting on page 624.

REFERENCES Ackefors, H. (1969). Ecological zooplankton investigations in the Baltic proper 1963-1965. Institute of Marine Research, Lysekil, Series Biology, Report No. 18, 1-137.

Adams, J. R. (1969). Ecological investigations around some thermal power stations in California tidal waters. Chmapeake Science, 10, 145-154. Alcock, A. and Anderson, A. R. (1894). An account of a recent collection of deep sea Crustacea from the Bay of Bengal and Laccadive Sea. Natural history notes from H.M. Indian Marine Survey steamer “Investigator”, Cmdr. C. F. Oldham, R.N., commanding. Journal of the Aeiatic Society of Bengal, 63, 141-185.

Alexandrowicz, J. S. (1955). Innervation of the heart of Praunus jZexuoaw, (Mysidacea). Journal of the Marine Biological Association of the United Kingdom, 34, 47-54. Alldredge, A. L. and King, J. M. (1977). Distribution, abundance and substrate preferences of demersal reef zooplankton a t Lizard Island Lagoon, Great Barrier Reef. Marine Biology, 41, 317-333. Almeida Prado, M. S. de (1966). Notes on the developmental stages ofSch&tomys& spiritus (Norman, 1860). Anais da Academia Braaileira de Cienciaa, 38, 349-353.

Almeida Prado, M. S. de (1972). Mysidacea (Crustacea) da regiao lagunar de Cananeia. Instituto OceanograJco da Universidad de Sao Paulo, 1-86. Almeida Prado, M. S. de (1973). Distribution of Mysidacea (Crustacea) in the Cananeia region. Bolitim. Zoologico e Biologic0 Marina, 30, 395-417. Almeida Prado, M. S. de (1974). Sistematica dos Mysidacea (Crustacea) na regiao de Cananeia. Boletim do Instituto OceanograJco Sao Paulo, 23, 47-87. Amaratunga, T. and Corey, S. (1975). Life history of Mysis stenokpk Smith (Crustacea, Mysidacea). Canadian Journal of Zoology, 53, 942-952. Amin, 0. M. (1978). On the crustacean hosts of larval acanthocephalan and cestode parasites in southwestern Lake Michigan. Journal of Parasitology, 64, 842-845.

Anonymous (1920). Report of Council for 1919. Scottish Marine Biological Association Annual Report, 3-5. Anonymous (1957). “Plymouth Marine Fauna” (Third Edn), 457 pp. The Marine Biological Association of the United Kingdom. Apstein, C. (1906). Lebensgeschichte von My& mixta der Ostsee. Wissenschaftliche Meeresuntersuchungen, 9, 241-260. Ariani, A. P. (1967). Observazione su Misidacei della costa Adriatica pugliese. Annuario dell’ Instituto e M w e o de Zoologia dell’ Universita d i Napoli, 18, 1-38.

Ariani, A. P. and Spagnuolo, G. (1976). Ricerche sulla misidofauna del Parco di Santa Maria di Castellabate (Salerno) con descrizione di una nuova specie di Siriella. Bolletino della Societa dei Naturalisti in Napoli, 84 (1975), 441481.

REFERENCES

265

Armitage, M. E., Raymont, J. E . G. and Morris, R. J. (1977). The effects of a pure carbohydrate diet on ‘the amino acid composition of Neomysis integer. In “A voyage of Discovery” (Ed. M. Angel), pp 471-481. Supplement t o Deep Sea Research, Pergamon Press, Oxford. Armitage, M. E., Raymont, J. E. G. and Morris, R. J. (1978).The effects of a pure protein diet on the amino acid composition of Neomysis integer (Leach). Journal of Experimental Marine Biology and Ecology, 35, 147-163. Amtz, W. E. (1971). Biomasse und Produktion des Makrobenthos in den tieferen Teilen der Kieler Bucht in Jahr 1968. Kieler Meeresforschung, 27, 36-72. Amtz, W. E . (197th). The food of adult cod (cladus morhua L.) in the Western Baltic. Meeresforschung, Reports on Marine Research, 26, 60-69. Amtz, W. E. (1978b). Predation on benthos by flounders, Platichthys jZesee (L.) in the deeper parts of Kiel Bay. Meeresforschung, Reports on Marine Research, 26, 70-78. &on, W. (1962). The distribution of animals in the eastern north Pacific and its relationship to physical and chemical conditions. Journal of the Fisheries Research Board of Canada, 19, 271-314. Austin, J. (1976). Preliminary observations on the mysid fauna of the N. shore of Belfast Lough. Irish Naturalist Journal, 18, 27&277. Baan, S. M: van der and Holthuis, L. B. (1971). Seasonal occurrence of Mysidacea in the surface plankton of the southern North Sea near the “Texel” lightship. Netherlands Journal of Sea Research, 5, 227-239. Bacescu, M. (1935). Metamysis strauchi (Czern.) Sars, Katamysis warpachowskyi Sars et Paramysis helleri (Sars), Mysidaces nouveaux pour la f a m e de la Roumanie. Annabs scientifiques de 1’ Universitd de Jassy, 21, 468-485. Bacescu, M. (19364. Prezenta Mysidelor in portiunea olteneasca a Dunarii. Revista stiinti$ca ‘‘ V . Adamachi”, Iasi, 23, 43-45. Bacescu, M . (1936b). Hemimysis lamornae sb. 8p. reducta et Hemimysis anomala dans les eaux Roumaines de la Mer Noire. Annales scienti3ques de 1’ Universitd de Jassy, 23, 70-93. Bacescu, M. (1938). Les mysidaces des eaux Roumaines. Annales scientiflqw de I’Universitd de Jassy, 26, 451-801. Bacescu, M. (1941). Les mysidaces des eaux MBditerraneennes de la France (sp6cialement de Banyuls) et des eaux de Monaco. Bulbtin de Z’lnstitut Ockanographique, Monaco, 795, 1-46. Bacescu, M. (1949). Donn6es sur la faune cacinologique de la Mer Noire de long de la cBte Bulgare. Travaux de la Station Biologique Maritime de Varna, Bulgaria, 14, 1-24. Bacescu, M. (1954).Fauna Republicii Populare Romine. Crustacea. IV Mysidacea. Academia Republicii Populare Romine, 1-126. Bacescu, M. (1961). Taphromysis bowmani, n.sp., a new brackish water mysid from Florida. Bulletin of Marine Science of the OuVand Caribbean, 11,517-524. Bacescu, M. (1966). Contribution a 1’6tude du genre Leptomysis et description de trois taxons nouveaux : Leptomysis peresi n.sp., L. apiops bayulensis n.ssp. et L. burgii a s p . Vie et Milieu. 17, 121-142. Bacescu, M. (1967). Further mysids from the Pacific Ocean collected during the X I t h cruise of R/V “Anton Bruun”, 1965. Revue R o u w i n e de Biologie, Sd& de Zoologie, 12, 147-159. Bacescu, M. (196th).Afromysis guinenais n.sp. and Brasilomysis castroi n.g., n.sp. from the waters of the tropical Atlantic. Revue Roumaine de Biologie, Sdrie de Zoologie, 13, 75-86.

266

THE BIOLOGY OF MYSIDS

Bacescu, M. (1968b). Heteromysini nouveaux des eaux Cubaines: trois esphces nouvelles de Heteromysis et ’ Heteromysoides spongicola n.g., n.sp. Revue Roumaine de Biologie, Sdrie de Zoologie, 13, 221-237. Bacescu, M. (196%). Etude des quelques Leptomysini (Crustacea Mysidacea) des eaux du Bresil et de Cuba; description d’un genre et de cinq autres taxons nouveaux. Annali det Muaeo civic0 di storia naturale Giacomo Doria, Genova, 77, 231-249. Bacescu, M. (1968d). Contributions to the knowledge of the Gastrosaccinae psammobionte of the tropical America, with the description of a new genus (Bourrnuniella n.g.) and three new species of its frame. Travaux du M w & d’histoire naturelle “Gr. Antipa”, 8, 355-373. Bacescu, M. (1969). Contribution it la connaissance du genre Metamy8idopsi.s W. Tattersall 1951. M . Swifti n.sp.-M. Mexicana n.n., confondues avec M . munda Zimmer. Revue Roumaine de Biologie, Sdrie de Zoologie, 14, 349357. Bacescu, M. ( 1970a).Contributions it 1’Btudemorphobcologique des Gastrosaccinae (Crustacea, Mysidacea) du versant est de I’Atlantique et de la MBditerranBe. Description de G. mediterraneus n.sp., c f . olivae n.sp. et a. roscoffenab n.sp. Revue Roumaine de Biologie, Sdrie de Zoologie, 15, 217-234. Bacescu, M. (1970b). New spongicolous Heteromysis of the Caribbean Sea (H. gomezi nap. and H. mariani n.sp.). Revue Roumaine de Biologie, Skrie de Zoologie, 15, 11-16. Bacescu, M. (1971a). Mysimenzies hadalis g.n., sp.n., a benthic mysid of the Peru Trench, found during cruise XI/1965 of R/V Anton Bruun (U.S.A.). Revue Roumaine de Biologie, Sdrie de Zoologie, 16, 3-8. Bacescu, M. (1971b). Contributions to the mysid Crustacea from the Peru-Chile Trench (Pacific Ocean). Anton Bruun Report, No. 7, 3-24. Bacescu, M. (1973a.).A new case of commensalism in the Red Sea: the mysid Idiomysis tsurnamali n.sp. with the Coelenterata Megalactis and Caasiopea. Revue Roumaine de Biologie, Sdrie de Zoologie, 17, 3-7. Bacescu, M. (197313).Anisomysis levi n.sp. from the Red Sea and the dichotomic key of the species belonging to the genus with description of a new taxon, Paranisorngsis n.sg. Revue Roumaine de Biologie, Sdrie de Zoologie, 18,173-180. New mysids from the littoral East African waters: Haplostylus Bacescu, M. (1973~). estafricana n.sp. and Animsomysis ijimai e&ajricana n.ssp. Revue R o u m i n e de Biologie, Sdrie de Zoologie, 18, 317-324. Bacescu, M. (1973d). Contribution it la connaissance des mysides benthiques de la Mer Rouge. Rapport et proces-verbaux des rdunions. Commksion internationale pour I’explcration scienti$que de la mer Mkditerrande, 21, 643-646. Bacescu, M. (1975). Contributions to the knowledge of the mysid (Crustacea) from the Tanzanian waters. University Science Journal, University of Dar e8 Salaam, 1, 39-61. Bacescu, M. (1976). Contribution B la connaissance des mysidaces (CrustacBs) de la CBte Lybienne, avec la description de deux nouvelles espbces, Neoheteromy& mulbri n.sg., n.sp. et Heteromysis lybiana n.sp. Revue R o u m i n e de Biologie, Sdrie de Zoologie, 21, 85-91. Bacescu, M. and Mayer, R. (1961). MalacostracBs (MysidaCea, Euphausiacee, Decapoda, Stomatopoda) du plancton diurne da la MBditerranBe etude bashe sur la materiel du Lamont Geological Observatory, Washington. Rapport et proces-verbaux des rdunions. Commission internationale pour l’exploration scienti$que de la mer Mdditerranb, 16, 183-202.

REFERENCES

267

Bacescu, M. and Muradian, Z. (1977). Contribution b la connaissance des peracarides des eaux du nord-est de Lybye. Rapport et prom-verbaw dea rduniona. Commkaion internationale pour l’exploration acienti,fique de la mer M&itewan&, 24, 111-112.

Bacescu, M. and Orghidan, T. (1971). Antromy&a c u b a n k n.sp. et Spelaeomysis nuniezi n.sp., Mysis cavernicoles nouvelles de Cuba. Revue Roumaim de Bwlogie, Sdrie de Zoologie, 16, 225-231. Bacescu, M. and Orghidan, T. N. (1977). New contribution to the knowledge of troglobian mysids of Cuba: AntromyG juberthiei n.sp. I n “Resultats des Exp6ditions Biospelogiques Cubano-Roumainesh Cuba”, Part 3, pp. 263-265. Editura Academiei Republicii Socialiste Romania, Bucuresti. Bacescu, M. and Schiecke, U. (1974). Qastroaaccw,magnilobatw, n.sp. and Erythrope peterdohrni n.sp. (Mysidacea)-new surprises from the Mediterranean benthos. Cmtaceana, 27, 113-118. Bacesu, M. and Vasilescu, E. (1973). New benthic mysids from the littoral waters of Kenya: Mysidopaia kenyana n.sp. and Nouvelia ndalenG naornbaaae n.g., nap. Revue Roumine de Biologk, Sdrie de Zoologie, 18, 249-256. Bagge, P. and nus, E. (1973). Interannual changes of the soft bottom fauna at some permanent Finnish stations in 1967-1970. AnmZea BiologGuea, 28 (1971), 78-85.

Bailey, R. M., Fitch, J. E., Herald, E. S., Lachner, E. A., Lindsey, C. C., Robins, C. R. and Scott, W. B. (1970). A list of common and scientific names of fishes from the United States and Canada. Amrican Fisheries Society, Special Publication, No. 6., 1-149. gainbridge, R. and Waterman, T. H. (1957). Polarized light and the orientation of two marine Crustacea. Journal of Experimental Biology, 34, 342-364. Bainbridge, R. and Waterman, T. H. (1958). Turbidity and polarized light orientation of the crustacean Mysidium. Journal of Experimental Biology, 35, 48 7-493.

Bainbridge, V. (1960). “he plankton of inshore water off Freetown, Sierra Leone. Colonial Ofice,Fi8hery Publication, No. 13, 1-48. Balasubrahmanyan,K. (1964). Mysids from the Bay of Bengal. I n “Some Aspects of Crustacea”, pp. 12-19. Marine Biological Station of Porto Novo, India. Balasubrahmanyan,K. and Prince, M. J. (1976). Pro&jua ovatw, Pillai (Isopoda) parasitic on a new host. Current Science, 45, p. 603. Bamstedt, U. (1978). Studies on the deep-waterpelagic communityof Korsfjorden, Western Norway. Seasonal variation in weight and biochemical composition ofChiridiusarmatw,(Copepoda),Boreomyabarctica (Mysidacea),and Eukrohnia hamta (Chaetognatha) in relation to their biology. Sarah, 61, 145-154. Banner, A. H. (1948). A taxonomic study of the Mysidacea and Euphausiacea (Crustacea) of the North Eastern Pacific. Part 11. Mysidacea, from tribe Mysini through subfamily Mysidellinae. Tranaactiona of the Royal C a d i a n In&&&, 27, 65-125. Banner, A. H. (1953). On the new genus and species of m p i d from Southern Louisiana. Tuhne Studies in Zoology, 1, 3-8. Banner, A. H. (1954a). A supplement to W. M. Tattersall’s Review of the Mysidacea of the United States National Museum. Proceedings of the United States National Museum, 103, 575-583. Banner, A. H. (1954b). Some “schizopod” crustaceans from the deeper water off California. A l h n Hancock Foulzdation Publicationa, Occasional Paper, No. 13, 1-48.

268

THE BIOLOGY OF MYSIDS

Barker, J. M. (1973). Changes in the total energy content of My sis relicta Loven during its vertical migration. Probeedings of the 16th Conference on Great Lakes Research, 1-6. Barnes, R . S. K., Dorey, A. E. and Little, C. (1971). An ecological study of a pool subject to varying salinity (Swanpool, Falmouth). Journal of Animal Ecology, 40, 709-734. Bary, B. M. (1956). Notes on ecology, systematics and development of some Mysidacea and Euphausiacea (Crustacea) from New Zealand. Pacific Science, 10, 431-467.

Beanland, F. L. (1940). Sand and mud communities in the Dovey Estuary. Journal of the Marine Biological Association of the United Kingdom, 24, 589-611.

Beaudouin, J. (1977). Euphausiacits et mysidacb du Golfe du Gascogne (Plateau Continental) en 1971. Annales Biologiques, 32 (1975), 67-69. Beck, J. T. (1977). Reproduct,ion of the estuarine mysid T’aphromysis bow~nuni (Crustacea : Malacostraca) in fresh water. Marine Biology, 42, 253-257. Bedard, J. (1969). Feeding of the least, crested and parakeet auklets around St Lawrence Island, Alaska. Canadian Journal of Zoology, 47, 1025-1050. Beeton, A. M. (1959). Photoreception in the opossum shrimp, My sis relicta Loven. Biological Bulletin, Marine Biological Laboratory, Woods Hole, Mass., 116, 204-216.

Beeton, A. M. (1960). The vertical migration of My sis relicta in Lakes Huron and Michigan. Journal of the Fisheries Research Board of Canada, 17, 517-539. Beeton, A. M. (1969). Changes in the environment and biota of the Great Lakes. I n “Eutrophication: causes, consequences, correctives”, pp. 150-187. National Academy of Sciences, Washington, D.C. Beeton, A. M. and Clarke, W. D. (1973). Mysid bibliography. Center for &eat Lakes Studies, the University of Wisconsin-Milwaukee,Special Report, No. 16, 1-145.

Beeton, A. M. and Edmondson, W. T. (1972). The eutrophication problem. Journal of the Fisheries Research Board of Canada, 29, 673-682. Bell, J. D., Burchmore, J. J. and Pollard, D. A. (1978). Feeding ecology of a scorpaenid fish, the fortescue Centropogon australis from a Posidonia seagrass habitat in New South Wales. Australian J o z x a l of Marine and Freshwater Research, 29, 175-185. Bellan-Santini, D. (1966). Contribution B 1’6tude du peuplement des cavitbs sciaphiles de l’encorbellement a Lithophyllum tortuosum dans la region Marseillaise. Recueil des Travaux de la Station Marine d’Endouww, 40, 151-157.

Belloc, G. (1959). Catalogue des types de mysidaces du Mus6e Oceanographique de Monaco. Bulletin de l’lnstitut Ocdanographique de Monaco, No. 1160, 1-6. Belman, B. W. and Childress, J. J. (1976). Circulatory adaptations to the oxygen minimum layer in the bathypelagic mysid Gnathophawria ingens. Biological Bulletin, Marine Biological Laboratory, Wood’s Hole, Mass., 150, 15-37. Belyaev, G. M. (1966). “Hadal bottom fauna of the world ocean”, 199 pp. Israel Program for Scientific Translations, Jerusalem, 1972. Belyayeva, V. N., Pirogovskiy, M. I. and Polyaninova, A. A. (1972). Distribution and food of beluga ( H M O huso (L)) underyearlings in the North Caspian. Voprosy ikhtiologii (English edn), 12, 87-94. Ben’ko, K. E. (1962). Some quantitative studies of the food of mysids. Trudy Karadahs’koyi nauchnoyi stantsiyi imeni T . I . Vyazems’koho, 18, 37-43.

REFERENCES

269

Berrill, M. (1969). The embryonic behaviour of the mysid shrimp, Mysis relicta. Canadian Journal of Zoology, 47, 1217-1221. Berrill, M. (1971). The embryonic development of the avoidance reflex of Neomysis amricana and Praunus flexuosw (Crustacea : Mysidacea). Animal Behaviour, 19, 707-713. Beyer, F. (1958). A new, bottom living Trachymedwa from the Oslofjord. Nytt Magasin for Zoologi, 6, 121-143. Biesinger, K. E., Lemke, A. E., Smith, W. E. and Tyo, R. M. (1976). Comparative toxicity of polyelectrolytes to selected aquatic animals. Journal of the Water Pollution Control Federation, Washington, 48, 183-187. Birstein, J. A. and Tchindonova, J. a. (1958). The deep sea mysids of the northwest Pacific Ocean. Trudy Inatituta okeanologii, Akadem/iya nauk SSSR, 27,25a355. Birstein, J. A. and Tchindonova, J. G. (1962). Mysidacea collected by the Soviet Antarctic Expedition on the M/V “Ob”. Biological Reports of the Soviet Antarctic Expedition (1955-1958),1, 58-68. Birstein, J. A. and Tchindonova, J. G. (1970). New mysids (Crustacea, Mysidacea) from the Kurile-Kamchatka Trench. Trudy Instituta okeanologii, Akademiya nauk SSSR, 86, 277-291. Blegvad, H. (1922). On the biology of some Danish gammarids and mysids. Report of the Danish Biological Station to the Board of Agriculture, 28, 1-103. Boesch, D. F. and Diaz, R. J. (1974). New records of peracarid crustaceans from oligohaline waters of the Chesapeake Bay. Chesapeake Science, 15, 56-59. Boesch, D. F., Diaz, R. J. and Virnstein, R. W. (1976). Effects of tropical storm Agnes on soft-bottom macrobenthic communities of the James and York estuaries and the lower Chesapeake Bay. Chesapeake Science, 17, 246-259. Bordovskiy, 0. K., Shirinskiy, V. G. and Akhmet’yeva, Ye. A. (1976). Isotopic composition of carbon in the lipids of zooplankton from the Kuril-Kamchatka area of the Pacific Ocean. Oceanology, 16, 568-570. Borgmann, U. (1978). The effect of temperature and body size on electron transport system activity in freshwater zooplankton. Canadian Journal of Zoology, 56, 634-642. Boroditch, N. D. (1978)..TheCaspian Peracarida (Crustacea) in the Saratov water reservoir. Zoologichesky Zhurnal, 57, 783-785. Boroditch, N. D. and Havlena, F. K. (1971). Mysidacea in the aft bay of the Volzhsky hydro-electric power station and the Saratov water reservoir. Zoologichesky Zhurnal, 50, 1101-1104. Boroditch, N. D. and Havlena, F. K. (1973). The biology of mysids acclimatized in the reservoirs of the Volga River. Hydrobiologia, 42, 627-539. Boschma, H. (1949). Ellobiopsidae. “Discowery” Reports, 25, 281-314. Boschma, H. (1959). Ellobiopsidae from tropical West Africa. Atlantide Report, 5, 145-175. Bossanyi, J. (1957). A preliminary survey of the small natant fauna in the vicinity of the sea floor off Blyth, Northumberland. Journal of Animal Ecology, 26, 353-368. Bowers, J. A. and Grossnickle, N. E. (1978). The herbivorous habits of My& relicta in Lake Michigan. Limnology and Oceanography, 23, 767-776. Bowman, T. E. (1957). A new species of M y d o p s i s (Crustacea: Mysidacea) from the south eastern coast of the United States. Proceedings of the United States National Muaeum, 107 (3378), 1-7.

270

THE BIOLOGY OF MYSIDS

Bowman, T. E. (1964). Mysidopsis alrnyra, a new estuarine mysid crustacean from Louisiana and Florida. Tulane Studies i n Zoology, 18, 15-18. Bowman, T. E. (1973). Two new American species of Spelaeomysis (Crustacea: Mysidacea) from a Mexican cave and land crab burrows. Association for Mexican Cave Studies, No. 5, 13-20. Bowman, T. E. (1976). Stygiomysis major, a new troglobitic mysid from Jamaica, and extension of the range of S. holthuisi, t o Puerto Rico (Crustacea: Mysidacea : Stygiomysidae). International Journal of Speleology, 8, 365373. Bowman, T. E. (1977a). Mysidacea. I n “Biota Acuatica de Sudamerica Austral” (Ed. S . H. Hurlbert), pp. 149-150. San Diego State University, California. Bowman, T. E. (197713).Areview of the genus Antromysis (Crustacea: Mysidacea), including new species from Jamaica and Oaxaca, Mexico, and a redescription and new records for A . cenotensis. Association for Mexican Cave Studies. No. 6, 27-38. Boyden, C. R., Crothers, J. H., Little, C. and Mettam, C. (1977). The intertidal invertebrate fauna of the Severn Estuary. Field Studies, 4, 477-554. Boysen, H. 0. (1976). Regionale und saisonale Unterschiede in der Besiedlungsdichte von Mysidaceen und Dekapodenlarven im Hyperbenthos der Kieler Bucht. Meeresforschung, Reports on Marine Research, 25, 54-63. Brattegard, T. (1969). Marine biological investigations in the Bahamas. 10. Mysidacea from shallow water in the Bahamas and southern Florida. Part I. Sarsia, 39, 17-106. Brattegard, T. (1970s). Marine biological investigations in the Bahamas. 11. Mysidacea from shallow water in the Bahamas and southern Florida. Part 11. Sarsia, 41, 1-36. Brattegard, T. (1970b). Mysidacea from shallow water in the Caribbean Sea. Sarsia, 43, 111-154. Brattegard, T. (1973). Mysidacea from shallow water on the Caribbean coast of Colombia. Sarsia, 54, 1-66. Brattegard, T. (19740). Additional Mysidacea from shallow water on the Caribbean coast of Colombia. Sarsia, 57, 47-86. Brattegard, T. (1974b). Mysidacea from shallow water on the Caribbean coast of Panama. Sarsia, 57, 87-108. Brattegard, T. (1975). Studies on the fauna of Curacao and other Caribbean Islands: No. 157. Shallow-water Mysidacea from the Lesser Antilles and other Caribbean regions. Uitgaven van de Natuurwetenschappelijke studiekring voor Surinam en dr Nederlandse Antillen, 83, 102-115. Brattegard, T. (1977). Three species of Mysidacea (Crustacea) from Surinam. Zoologische Mededelingen, 50, 283-293. Brattegard, T. and Vader, M. (1972). A collection of Peracarida from More and Romsdal, northwestern Norway. Sarsia, 49, 33-40. Brattelid, T. E. and Matthews, J. B. L. (1978). Studies on the deep-water pelagic community of Korsfjorden, western Norway. The dry weight and caloric content of Euchaeta norvegica (Copepoda), Boreomysis arctica (Mysidacea), and Meganyctiphanes norvegica (Euphausiacea). Sarsia, 63, 203-21 1. Broch, H. (1927). Untersuchungen iiber die Marine Bodenfauna bei Lindesness im Juni 1926. Avhandlinger utgitt av det Norske Videnskap-Akademi i 0810, Matematisk-NaturvidenskapeligKlasse, ( 5 ) , 1-32. Brown, A. C. (1964). Food relationships on the intertidal sandy beaches of the Cape Peninsula. South African Journal of Science, 60, 35-41.

REFERENCES

271

Brown, A. C. (1971). The ecology of sandy beaches of the Cape Peninsula, South Africa. Part I. General introduction. Transactions of the Royal Society of South Africa, 39, 247-280. Brown, A. C. and Talbot, M. S. (1972). The ecology of the sandy beaches of the Cape Peninsula, South Africa. Part 3. A study of Gaatrosaccue p s a m d y t e e Tattersall (Crustacea : Mysidacea). Transaction8 of the Royal Society of South Africa, 40, 309-333. Brownell, W. N. (1970). Comparison of Mysis relicta and Pontoporeia a$inis as possible intermediate hosts for the acanthocephalan Echinorhymhue salmonis. Journal of the Fhheries Research Board of Canada, 27, 1864-1866. Brunel, P. (1959). Le zooplankton de la Baie des Chaleurs en 1955: distribution horizontale quantitative et correlations hydroclimatiques. Department des Pbheries, Quebec, Contribution, No. 73, 1-65. Brunel, P. (1960). Artificial key to the Mysidacea of the Canadian Atlantic Continental Shelf. Canadian Journal of Zoology, 38, 851-855. Brunel, P. (1961). Liste taxonomique des invertebres marins des parages de la Gaspesie identifies au 3 Aoilt 1959. Station de Biologie Marine de GrandeRivihe, Cahhr8 d ’ I n f o r m d w n , No. 7, 1-9. Brunel, P. (1963a). Inventaire taxonomique des invertebr6s marins du Golfe Saint-Laurent. Station de Biologie Marine de Grande-RiViBre, Rapport Annuel, 1962, 81-89. Brunel, P. (196313).Variations journalibres et saisonnihres de l’alimentation de la morue au large de Grande Rivi6re. Station de Biologie Marine de GrandeR i d r e , Rapport Annuel 1962,101-117. Brunel, P. (1965). Inventaire taxonomique des invertebres benthiques marins du Golfe Saint-Laurent.Station de Biologie Marine de Grande-RiviBre, Rapport Annuel, 1964, 1-2. Brunel, P. (1970). Catalogue d’invertebr6s benthiques du Golfe Saint-Laurent recueilles de 1951 It 1966 par la station de Biologie Marine de Grande-Rivibre. Travaux sur l w Pt!chel.iw duQuebec, No. 32,l-54. Bruun, A. F. (1957).Deep sea and abyssal depths. I n “Treatise on Marine Ecology and Palaeoecology” (Ed. J. W. Hedgpeth), 1, pp. 641-672. Geological Society of America Memoir 67. Burton, J. D. and Massie, K. S. (1971). The occurrence of tantalum in some marine organisms. Journal of the Marine Biological Association of the United Kinqdm, 51, 679-683. Calman, W. T. (1932). A cave-dwelling crustacean of the family Mysidae from the island of Lanzarote. Annals and Magazine of Natural History, Ser. 10, 10, 127-131. Camp, D. K., Whiting, N. H. and Martin, R. E. (1977).Nearshore marine ecology at Hutchinson Island, Florida: 1971-1974. IV. Arthropods. Florida Marine Research Publications, No. 25, 1-63. Cannon, H. G. and Manton, S. M. (19274. On the feeding mechanism of a mysid crustacean: Hemimysis lanwrnae. Transactiona of the Royal Society of Edinburgh, 55, 219-253. Cannon, H. G. and Manton, S. M. (1927b). Notes on the segmental excretory organs of Crustace-I-IV. Journal of the Linnaean Society, Zoology, 36, 439456. Caroli, E. (1937). Stygiomysis hydruntina n.g., n.sp., Mysidaceo carvernicolo di Terra d’otranto, rappresentante di m a nuova famiglia. Bolbtino Zoologia, Torino, 8, 219-227.

272

THE BIOLOGY OF MYSIDS

Carpenter, G. F., Mansey, E. L. and Watson, N. H. E. (1974). Abundance and life history of Mysis relicta in the St Lawrence Great Lakes. Journal of the Fisheries Research Board of Canada, 31, 319-325. Casanova, J.-P. (1968). Sur un mysidace pelagique peu connu: Euchaetomeropeia merolepis Illig, 1908. Crustaceam, 15, 153-160. Casanova, J.-P. (1970). Essai de classement bathymetrique des formes zooplanctoniques en MediterranBe. Revue des Travaux, Institut dea Hches rnaritirnes, 31, 45-58. Casanova, J.-P. (1977). La faune pelagique profonde (zooplancton et micronecton) de la province Atlanto-Mediterranbenne.Aspects taxonomique, biologique et zoogeogmphique. These presentee B l’Universit6 de Provence (Aix-Marseille I) pour obtenir le grade de Docteur es-Sciences Naturelles, 1-465. Caspers, H. (1957). Black Sea and Sea of Azov. I n “Treatise on Marine E c o l o g ~ end Palaeoecology” (Ed. J. W. Hedgpeth), 1, pp. 801-889. Geological Society of America, Memoir 67. Cerruti, A. (1921). Ulteriori notizie biologiche riguardanti specialmente il period0 di maturita sessuale degli animali del Golf0 di Napoli. Pubblicazioni della Stazione zoologica di Napoli, 3, 229-240. Champalbert, G. and Macquart-Moulin, C. (1970). Les peracarides de l’hyponeuston nocturne du Golfe de Marseille. Cahiers de Biologie marine, 11, 1-29.

Charniaux-Cotton, H., Zerbib, C. and Meusy, J. J. (1966). Monographie de la glande androgene des crustacbs superieurs. Crushceana, 10, 113-136. Chicewicz, M. (1952). A study of colour change in Praunus flexuosm Miill. Bulletin international de I’Academiepolonaise des sciences et des lettres, Zoologie, Ser. B, 11, 371-384. Childress, J. J. (1968). Oxygen minimum layer: vertical distribution and respiration of the mysid Qmthophawia ingens. Science, 160, 1242-1243. Childress, J. J. (19714. Respiratory adaptations to the oxygen minimum layer in the bathypelagic mysid Gnathophawia ingens. Biological Bulletin, Marine Biological Laboratory Wood’s Hole, Maas., 141, 109-121. Childress, J. J. (1971b). Respiratory rate and depth of occurrence of midwater animals. Limnology and Oceanography, 16, 10P106. Childress, J. J. (1975). The respiratory rates of midwater crustaceans as a function of depth of occurrence and relation to the oxygen minimum layer off southern California. Deep-sea Research, 50A, 787-799. Childress, J. J. and Nygaard, M. (1974). Chemical composition and buoyancy of midwater crustaceans as function of depth of occurrence off southern California. Marine Biology, 27, 225-238. Childress, J. J. and Price, M. H. (1978). Growth rate of the bathypelagic crustacean Gnathophausia ingens (Mysidacea: Lophogastridae). 1 Dimensional growth and population structure. Marine Biology, 50, 47-62. Childress, J. J., Barnes, A. T., Quetin, L. B. and Robison, B. H. (1978). Thermally protecting cod ends for the recovery of living deep-sea animals. Deep-sea Research, 25, 419-422. Chin, P.(1971). The effect of temperature on respiratory metabolism of the mysid, Neomysis awatschensis (Brandt ). Publications of the Marine Laboratory of Pusan Fisheries College, 4, 9-18. Chin, P. (1972). The effect of salinity on respiratory metabolism of the mysid, Neomysis awatschensis (Brandt). Publications of the Marine Laboratory of P w a n Fishelie-9 College, 5, 31-36.

REFERENCES

273

Chin, P. (1974). Acclimation and metabolic attitude of Neomysis awatschensis. Publication8 of the Marine Laborat09 of Pusan Fisheries College, 7, 1-20. Chin, P. (1976). Nitrogen and phosphorus excretion in Neomysis awatschensis. 1. Effects of temperature and salinity. Publications of the Imtitute of Marine Science, National Fisheries University of Buaan, 9, 1-6. Christensen, M. S. (1978). Trophic relationships in juveniles of three species of sparid fishes in the South African marine littoral. Fishery Bulletin, National Oceanic Atmospheric Administration of the United,States, 76, 389-401. Chun, C. (1896). Atlantis. Biologisohe Studien uber pelagische Tiefsee-Schizopoden. Zoologica, Stuttgart, 7, 1-262. Cinelli, F., Fresi, E., Mazzella, L., Pansini, M., Pronzato, R. and Svoboda, A. (1977). Distribution of benthic phyto- and zoocoenoses along a light gradient in a superficial marine cave. I n “Biology of Benthic Organisms” (Eds B. F. Keegan et al.) pp. 173-183. Pergamon Press, Oxford. Clarke, T. A. (1978). Die1 feeding patterns of 16 species of mesopelagic fishes from Hawaiian waters. Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 76, 495-513. Clarke, W. D. (1955). A n0w species of the genus Heteromysis (Crustacea, Mysidacea) from the Bahama Islands, commensal with a sea-anemone. American Museum Novitates, No. 1716, 1-13. Clarke, W. D. (1961a). A giant specimen of Cnathophausia ingens (Dohm, 1870) (Mysidacea)and remarks on the asymmetry of the paragnaths in the suborder Lophogastrida. Crustaceana, 2, 313-324. Clarke, W. D. (1961b). Proposal of a new name, Lepidomysis, for the preoccupied mysidacean generic name Lepidops Zimmer, 1927. Crustaceana, 2, 251252.

Clarke, W. D. (1962). The genus Gnathophausia (Mysidacea, Crustacea), its systematics and distribution in the Pacific Ocean. Doctoral Thesis, University of California, San Diego, 1-251. Clutter, R. I. (1965). Self-closing device for sampling plankton near the sea bottom. LimnoEogy and Oceanography, 10, 293-296. Clutter, R. I. (1967). Zonation of nearshore mysids. Ecology 48, 200-208. Clutter, R. I. (1969). The microdistribution and social behaviour of some pelagic mysid shrimps. Journal of Experimental Marine Biology and Ecology, 3, 125-155.

Clutter, R. I. and Theilacker, G. H. (1971). Ecological efficiency of a pelagic mysid shrimp; estimates from growth, energy budget and mortality studies. Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 69, 93-115. Colman, J. S. (1962). A note on deep plankton in the Bay of Biscay. Rapport et proceu-verbaux des rkunions. Conseil permanent international pour .!’exploration de la m r , 153, 207-210. Colman, J. S. and Segrove, F. (1955). The tidal plankton over Stoupe Beck Sands, Robin Hood’s Bay. Journal of Animal Ecology, 24, 445-462. Colosi, G. (1916). Nuova diagnosi e posizione sistematica di Lycomysis spinicauda Hansen. Minotore zoologico italiano, 27, 193-200. colosi, G. (1923). Alcune specie discusse di Misidacei. Bollettino della Societa dei Naturalisti in Napoli, 35, 191-195. Colosi, G. (1930). Remarques sur la distribution horizontale et verticale des mysides mediterran6ens. Bulletin de la Sociiti d’Ocianographie de France, 10, 983-985.

274

THE BIOLOGY OF MYSIDS

Connell, A. D. (1974). Mysidacea of the Mtentu River Estuary, Transkei, South Africa. Zoologica Africana, 9,. 147-159. Costa, H. R. Da. (1964). Notas sobre 0s Mysidacea da Costa brasileira. Boktim . d o Museu Nacional Brasil, New Series, Zoologia, No. 247, 1-9. CoutiBre, M. H. (1904). Sur quelques points de la morphologie des Schizopodes. Annals and Magazine of Natural History, Ser. 7 , 13, 156-159. CoutiBre, M. H. (1917). Crustacea schizopodes et dbcqodcs. Ccuxidme Expddition Antarctique Francaise (1908-1910), Command& par le Dr Jean Charcot, 1-8.

Cronin, E., Daiber, J. C. and Hulburt, E. M. (1962). Quantitative seasonal aspects of zooplankton in the Delaware River Estuary. Chesapeake Science, 3, 63-93. Crouau, Y. (1978). Organes sensoriels d’un mysidacb souterrain anophthalme, Antromysis juberthiei : Btude ultrastructurale des aesthetascs. Bulletin du Mushe d’histoire nature&, section zoologie, 352, 165-175. Curl, H. (1962). Standing crops of carbon, nitrogen and phosphorus and transfer between trophic levels in continental shelf waters south o f New York. Rapport et proces-verbaux des rdunions. Conseil pemnanent international pour l’exploration de la mer, 153, 183-189. Cussans, M. (1904). Gammarus. Liverpool Marine Biological Committee, Memoir 12, 1-47.

Czerniavsky, V. ( 1882). Monographia Mysidarum imprimis imperii. Rossici. Fascicle 11. Travaux de la SociBtB irnperkle de8 Natwalistes de St Petersbourg, 13, 1-85. Dadswell, M. J. (1975). Some notes on shoaling behaviour and growth of My& gaspensis (Mysidacea)in a small Newfoundland estuary. Canadian Journal of Zoology, 53, 374-377. Dahl, E. (1956). On the differentiation of the topography of the crustacean head. Acta Zoologica, 37, 123-192. Dahl, E. and Mecklenburg, C. V. (1969). The sensory papilla X-organ in Boreomysis Zellarctica (Kreyer) (Crustacea Malacostraca Mysidacea). Zei&schrijt forschung und mikroskopische Anatomie, 101, 88-97. Dakin, W.J. and Colefax, A. N. (1940). The plankton of the Australian coastal waters off New South Wales. Part I. Publications of the University of Sydney, Department of Zoology, Monograph 1, 1-215. Daniel, R . J. (1928). The abdominal muscular systems of Praunw, jlexuoau.9 (Muller). Lancmhire Sea Fisheries Laboratory Report for 1927, 36, 5-41. Daniel, R . J. (1931). Comparative study of the abdominal musculature in Malacostraca. Part I. The main ventral muscles of the typical abdominal segments. Lancashire Sea Fisheries Laboratory Geport for 1930, 39, 57-71. Daniel, R . J. (1932). Comparative study of the abdominal musculature in Malacostraca. Part 11. The superficial and main ventral muscles, dorsal muscles, and lateral muscles, and their continuations into the thorax. Lancashire Sea Fisheries Laboratory Report f o r 1931, 40, 46-107. Daniel, R . J. (1933). Comparative study of the abdominal musculature in Malacostraca. Part 111. The abdominal muscular systems of Lophogaster typicus M. Sars and Gnathophausia zoea Suhm, and their relationships with the musculatures of other Malacostraca. Lancmhire Sea Fisheries Laboratory Report for 1932, 41, 71-133. Darnell, R. M. (1958). Food habits of fishes and larger invertebrates of Lake Pontchartrain, Louisiana, an estuarine community. Publications of the Institute of Marine Science, University of Term, 5, 353-416.

REFERENCES

275

Darnell, R. M. (1961). Trophic spectrum of an estuarine community based on studies of Lake Pontchartrain, Louisiana. Ecology, 42, 553-568. Davis, C. C. (1966). A study of the hatching process in aquatic invertebrates. XXII. Multiple membrane shedding in Myaidium columbiae (Zimmer) (Crustacea: Mysidacea). Bulletin of Marine Science of the Qulf and Caribbean, 16, 124-131. Davis, C. C. (1968). Mechanisms of hatching in aquatic invertebrate eggs. Oceanography and Marine Biology, an Annual Review, 6 , 325-376. Day, H. J. (1967). The biology of Knysns Estuary, South Africa. I n “Estuaries” (Ed. G. H. Lauff), pp. 397-407. American Association for the Advancement of Science, Washington, D.C., Publication No. 83. Day, J. H., Field, J. G. and Montgomery, M. P. (1971). The use of numerical methods to determine the distribution of the benthic fauna acrosa the continental shelf of North Carolina. Journal of Animal Ecology, 40, 93-125. Debaisieux, P. (1947). Statocystes de Praunua et leurs poils sensoriels. A n d m de la Socidtd Royale de Zoologie de Belgique, 7 8 , 26-31. Debaisieux, P. (1949). Les poils sensoriels d’arthropodes et l’histologie nerveuse. I. Praunuafixuoaua Mull. et Crangon crangon L. Cellule, 52, 311-360. Dediu, I. I. (1966). Repartition et charact6ristique Bcologique des mysides des bassins des riviers Dniestr et Pruth. Revue Roumaim de Biologk-Zoologie, 11, 233-239. Delage, Y. (1883). Circulation et respiration chez les crustac6s schizopodes. Archives de Zoologie experimentale et gkndrale. Ser. 2, 1, 105-130. Delamare Deboutteville,. C. (1950).Une station nouvelle du mysidac6 Heteromyk a m r i c a n a Nouvel et le biotope de cette esp6ce. Vie et Milieu, 1, 81-82. Delamare Deboutteville, C. (1953a). La faune des eaux souterraines littorales des plages de Tunisie. Vie et Milieu, 4, 141-170. Delemare Deboutteville, C. (1953b). Description d’un appareil pour la captwe de la f a m e des eaux souterraines littorales sous la mer. Premiers resultats. Vie et Milieu, 4, 411-421. La faune des eaux souterraines littorales en Delamare Deboutteville, C. (1953~). Algerie. Vie et Milieu, 4, 470-503. Deniel, C. (1974). Regime alimentaire des jeunes turbots Scophthalmua nuzximu-3 L. de la classe 0 dans leur milieu naturel. Cahiera de Bwlogie marine, 15, 551-566. Depdolla, P. (1923).Ndvu.ng und Nahrungsewerb bei P r a u n ~ ~ e x ~ (Miiller). 8ua Biologiaches Zentralblatt, 43, 634-546. Dion, Y. and Nouvel, H. (1960). Mysidaces et euphausiac6s recoltes en Mediterran6e occidentale par le navire octhnographique “President-Theodore-Tissier” en 1949; presence de 1’EllobiopsidaeA d l o c y s t i a fagei Boschma en MBditerran6e et sur un hBte nouveau. Bulletin dea Travaux publib par la station d’tlquiculture et de Ptche de Castiglione, 10,1958-1959,ll-19. Dormaar, K,A. and Corey, S. (1973). Some aspects of osmoregulation in M y s k atenoleph (Crustacea, Mysidacea). Jmrnal of the Fiaherim Research Board of Canuda, 30, 1747-1749. Dormaar, K. A. and Corey, S. (1978). Some aspects of osmoregulation in My& relicta Loven (Mysidacea). Cruataceamz, 34, 90-93. Dragovich, A. (1969). Review of studies of tuna food in the Atlantic Ocean. United States Fiah and Wildlife Service, Special Scienti$c ReporbFiqheries, 593, 1-21.

276

THE BIOLOQY OF MYBIDS

Dragovich, A. (1971). The food of skipjack and yellowfin tunas in the Atlantic Ocean. Fishery Bulletin, National .Oceanic Atmospheric Administration of the United States, 68, 445-460. Drainville, G., Tiphane, M. and Brunel, P. (1963). CroisiAre oc6anographique dens le fjord du Saguenay 14-22 Juin, 1962. Cahiers de l’lnfomnation, Station Biologique Marine Urasde-Riviire, 17, 133-148. Dryer, W. R., Erkkila, L. F. and Tetzloff, C. L. (1965). Food of the lake trout in Lake Superior. Transactions of the American Fishrim Society, 94, 169176.

Dunbar, M. J. (1940). On the size distribution and breeding cycles of four marine planktonic animals from the Arctic. Journal of A n i m l Ecology, 9, 215-226. Dunbar, M. J. (1941). On the food of seals in the Canadian Eastern Arctic. Canadian Journal of Research, D, 19, 150-155. Dunbar, M. J. (1942). Marine macroplankton from the Canadian Eastern Arctic. I. Amphipoda and Schizopoda. Canadian Journal of Reaearch, D, 20, 33-46. Edwards, R. and Steele, J. H. (1968). The ecology of 0-Group plaice and common dabs a t Loch Ewe. I. Population and food. Journal of Experimental Marine Biology and Ecology, 2 , 215-238. Eggers, D. M., Bartoo, N. W., Rickard, N. A., Nelson, R. E., Wiasmar, R. C., Burgner, R. L. and Devol, A. H. (1978). The Lake Washington ecosystem: the perspective from the fish community production and forage base. Journal of.& Fisheries Research Board of Canada, 35, 1553-1571. Eguchi, E. and Waterman, T. H. (1966). Fine structure patterns in crustacean rhabdoms. I n “The Functional Organization of the Compound Eye”, pp. 105-124. Pergamon Press. Oxford and New York. Eguchi, E. and Waterman, T. H. (1973). Orthogonal microvillus pattern in the eighth rhabdomere of the rock crab Brapaw. Zeitschrift f fir Zellforschung und mikroskopische Amtomie, 137, 145-147. Ellertsen, B. (1977). A new apparatus for sampling surface fauna. Sarsia, 63, 113-114.

Elofsaon, R. (1965). The nauplius eye and frontal organs in Malacostraca (Crustacea). Sarsia, 19, 1-54. Elofsson, R. and Dahl, E. (1970). The optic neuropiles and chiasmata of Crustacea. Zeitachrift f fir Zellfmmhung und mikroskopische Anatomie, 107, 343-360. Elofsson, R. and Hallberg, E. (1973). Correlation of ultrastructure and chemical composition of crustacean chromatophore pigment. Journal of Ultrastructure Research, 44, 421-429. Elofsson, R. and Hallberg, E. (1977). Compound eyes of some deep-sea and fjord mysid crustaceans. A d a Zoobgica, 58, 169-177. Emery, A. R. (1968). Preliminary observations on coral reef plankton. himnology and Oceanography, 13, 293-303. Enbysk, B. J. and Linger, F. I. (1966). Mysid statoliths in shelf sediments off northwest North America. Journal of Sedimentary Petrology, 30, 839-840. Engel, W. A. V. and Tan, E.-C. (1965). Investigations of inner continental shelf waters off lower Chesapeake Bay. Part IV. The Copepods. CheeapeakeScience, 6, 183-189.

Enomoto, U. (1964). Studies on the food base in the Yellow and the Eaat China Seas. V. On the main crustaceans as food item of fish resources. Bulletin of the Japanese Society of Scientifcc Fisheries, 30, 216-226. Enright, J. T. (1962). Responses of an amphipod to pressure changes. Comparative Biochemistry and Physiology, 7 , 131-146.

REFERENOES

277

Evans, F. (1961). The planktonic Crustacea of the Petula trans-Atlantic Expedition. Proceedings of the ‘LinneanSociety, 172, 189-207. Fage, L. (1925). Lepidophthalmua serwatzu, Fage. Type nouveau de mysidac6 des eaux souterraines de Zanzibar. Biospeologica, Archivea de Zoologie experimental et gkn6rde, 63,826532. Fage, L. (1936). Sur un ellobiopside nouveau, Amallocystia faeciatwr, g. et sp. nov., parasite mysides bathypelagiques. Archives de Zoologie experimentale et gdndral, 78, 145-154. Fage, L. (1940). Sur le determinisme des caracthres sexuels secondaires des Lophogastrides (crustacBs-mysidac6s). Compte Rendu Hebdontadaire dea Seances de I’Academie des Sciences, 211, 335-337. Fage, L. (1941). Mysidacea. Lophogastrida I. Dana Reports, 4, (19), 1-52. Fage, L. (1942). Mysidacea. Lophogastrida 11. Dana Reports, 4 (23), 1-67. Fage, L. (1952). Etude Bcologique de la variation des carecthes differentiels des esphces et des populations. Annales de la Sociktd Royale Zoologique de Belgique,’ 83, 1-20.

Fege, L. and Monod, T. (1936). La faune marine du Jameo de Ague, lac souterraine de 1% de Lanzarote (Canaries). Archives de Zoologie experimentale et gkndrale, 78, 97-113.

Fager, E. W. and Clutter, R. I. (1968). Parameters of a natural population of a hypopelagic marine mysid, Metamysidopab elongata Holmes. Physiological ZOOZOgy741,267-267. Fager, E. y..,Flechsig, A. O., Ford, R. F., Clutter, R. I. and Ghelardi, R. J. (1966). Equipment for use in ecological studies using SCUBA. Limnology and Oceanography, 11, 503-509. Fain-Maurel, M.-A., Reger, J. F. and Cassier, P. (1975a). Le gamete mhle des schizopodes, et ses analogies avec celui des autres PBracarides. 1. Le spermatozoide. Journal of Ultrmtructure Reaearch, 51, 269-280. Fain-Maurel, M.-A., Reger, J. F. and Cassier, P. (1975b). Le gamete male des schizopodes, et ces analogies avec celui des autres PBracarides. 2. Particularit6 de la differenciation cellulaire au cows de la spermatogen8se. Journal of Ultraetructure Research, 51, 281-292. Field, J. G. (1971). A numerical analysis of changes in the soft-bottom fauna along a transect across False Bay, South Africa. Journal of Experimental Marine Biology and Ecology, 7 , 215-253. Fishelson, L. and Loya, Y . (1968). Preliminary observations on a population of Qmtroaaccua 8anctus (Van Beneden) (Mysidacea, Gastrosaccinae) on a Mediterranean sand beach of Israel. Crmtuceana, 15, 149-152. Fisher, L. R. and Kon, S. K. (1959). Vitamin A in the invertebrates. Biological Reviews, 34, 1-36. Fisher, L. R., Kon, S. K. and Thompson, S. Y. (1953). Vitamin A and carotenoids in some Mediterranean Crustacea with a note on the swarming of Meganyctip h n e a . Bulletin de Z’lnstitut Ockanographique de Monaco, No. 1021, 1-19.

Fisher, L. R., Kon, S. K. and Thompson, S. Y. (1955). Vitamin A and carotenoids in certain invertebrates. 111. Euphausiacea. Journal of the Marine Biological Association of the United Kingdom, 34, 81-100. Fleminger, A. and Clutter, R. I. (1965). Avoidance of towed nets by zooplankton. Limnology and Oceanography, 10, 96-104. Foul&, J. B. and Mann, K. H. (1978). Cellulose digestion in Mysie stenolepis and its ecological implications. Limnology and Oceanography, 23, 760-766.

278

THE BIOLOGY OF MYSIDS

Foulds, J. B. and Roff, J. C. (1976). Oxygen consumption during simulated vertical migration in Mysis relicta (Crustacea, Mysidacea). Canadian Journal Of Zoology, 54, 377-385. Fox, D. L., Oppenheimer, C. H. and Kittredge, J. S. (1953). Microfiltration in oceanographic research. Journal of Marine Research, 12, 233-243. Fox, H. M. (1952). Anal and oral intake of water by Crustacea. Journal of Experimental Biology, 29, 583-599. Foxon, G. E. H. (1940). The reactions of certain mysidR to stimulation by light and gravity. Journal of the Marine Biological Association of the United Kingdom. 24, 89-97. Franqueville C. (1971). Macroplancton profond (invertBbr6s) de la MBditerranBe nord-occidentale. Tethys, 3, 11-56. Fraser, J. H. (1950). List of rare exotic species found in the plankton by the Scottish research vessels in 1949. Annales Biologiques, 6, (1949), 95-99. Fraser, J. H. (1956). Scottish investigations from Aberdeen. Annales Biologiques, 11, (1944), 87. Fraser, J. $€. (1961). The oceanic and bathypelagic plankton of the northeast Atlantic and its possible significance to fisheries. Marine Research, 4, 1-48. Fraser, J. H. (1970). Scottish plankton collections 1969; zooplankton. Annubs Biologiques, 26 (1969), 112. Fraser, J. H. and Saville, A. (1949a). Macroplankton in the Faroe Channel, 1948. Annales Biologiques, 5 (1948), 29-30. Fraser, J. H. and Saville, A. (194913). Plankton distribution in Scottish and adjacent waters in 1948. Annales Biologiques, 5 (1948), 61-62. Freel, R. W. (1978). Oxygen affiity of the hemolymph of the mesopelagic mysidacean Cfnathophausia ingena. Journal of Experiwaental Zoology, 204, 267-273. Friedl, W. A. (1971a) Sonic scattering and its probable causes in two areas of Puget Sound. I n “Biological Sound Scattering in the Oceans” (Ed. G. B. Farquhar), pp. 527-548. Maury Center for Ocean Science, Department of the Navy, Washington, D.C. Friedl, W. A. (1971b). The relative sampling performance of 6- and 10-foot Isaacs-Kidd midwater trawls. Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 69, 427-432. Fujika, T., Yamamoto, T., Yamagi, I. and Shipematsu, T. (1969). Ash, iron, manganese content of marine plankton. Fisheries Research Board of Canada, Translation Series, No. 2557 (1973), 1-25. Fukai, R., Suzuki, H. and Watanabe, K. (1962). Strontium-90 in marine organisms during the period 1957-1961. Bulletin de I’Institut Ochpographique de Monaco, No. 1251, 1-16. Furnestin, M. L. (1959). Mysidaces du plancton marocain. Revues des Travaux, Institut de P b k mritimes, 23, 297-316. Furnestin, M. L. (1960). Zooplancton du Golfe du Lion et de la c6te orientale de Corse. Revues des Travaux, Institut de Ptches maritimes, 24, 153-252. Fiirst, M. (1965). Experiments on the transplantation of Mysia relicta into Swedish lakes. Report of the Institute of Freshwater Research, Drottningholm, 46, 79-89. Fiirst, M. (1967). Successful introducticns of glacial relicts as argument in a discussion of postglacial history. Report of the Institute of Freshwater Research, Drottningholm, 47, 113-117.

REFERENCES

279

Fiirst, M. and Nyman, L. (1969).Isoenzyme polymorphism in My& relic& Loven. Report of the Institute of Freshwater Research, Drottningholm, 49, 113117. Fuzessery, A. M. and Childress, J. J. (1975). Comparative chemosensitivity to amino acids and their role in the feeding activity of bathypelagic and littoral crustaceans. Biological Bulletin, Marine Biological Laboratory, Wood’s Hole, MWS., 149, 522-538. Gabe, M. (1952). Sur l’existence, chez quelques crustacbs malacostracBe, d’un organe comparable it la glande de la mue des insectes. C m p t e Rendua dea SeanceS de Z’Academie des Sciences, 237, 11 11-1 113. Gabe, M.(1956).Histologie comparBe de la glande de mue (organeY )des crustac6s maletcostrac6s. Ann& des Science0 Naturelles, Zoologie et Bwlogie Animale, 18, 145-152. Gabe, M. (1966).“Neurosecretion”, 872 pp. Pergamon Press, Oxford. Galt, J. H. and Whialer, H. C. (1970).Differentiation of flagellated spores in Thahsomyces ellobiopsid parasite of marine Crustaceae. Archiv f u r Mikrobiologie, 71, 295-303. Ganssle, D. (1966).Fishes and decapods of San Pablo and Suisun Bays. Californiu Piah and Game, Fishery Bulletin, 133, 64-94. Gaudy, R. and Guerin, J. P. (1979). Ecophysiologie compar6e des mysidac6s Hemimysia speluncola Ledoyer (cavernicole) et Leptomyaia lingvura G. 0. Sara (non cavernicole). Action de la temperature sur la croissance en Blevage. Journal of Experimentul Marine Biology and Ecology, 38, 101-119. Gauld, D.T. and Raju, R. C. S. (1976).Some measurements of the respiration of planktonic Crustacea. Bulletin of the Department of Marine Science, Univer8ity of Cochin, 7, 201-203. Geiger, S. R. (1969). Distribution and development of myside (Crustacea, (Mysidacea)from the Arctic Ocean and confluent seas. Bulletin of the Southern California Acwkmy of Sciences, 68, 103-111. Gelderd, C. (1907).Research on the digestive system of the Schizopoda. Anatomy, histology and physiology. Cellule, 25, 1-68. Genovese, S. (1956).Su due Misidacei dei laghi di Ganzirri e di Faro (Messina). Bolletino di Zoologia, pubbliuzto dall’ Unwne Zoologica Italians, 23, 177-196. Genovese, S. (1957). Analisi biometrica di una populazione di Siriella c l a d (Mysidacea, Siriellinae) vivente nel Lago di Faro. Bolletino d i Pam, Piscicoltura e Idrobiologia, 11, 246-263. Ghirardelli, E. (1973).Mer Adriatique. Rapport et procea-verbaw dea r6uniom. Cornmiasion internationale pour l’exploration de la w r , 21, 397-404. Gilson, G. (1909).Prodajw ostendensis n.sp. Etude monographique d’un epicaride parasite du Gaatrosaccua spinifer Goes. Bulletin ScientiAue de b France et de la Belgique, 43, 19-92. Gladfelter,W.B. (1979).Twilight migrations and foraging activities of the copper sweeper Pempheris schomburgki (Teleostei: Pempheridae). Marine Bwbgy, 50, 109-119. Goodbody, I. (1965). Continuous breeding in populations of two tropical crustaceans, Mysidium columbiae (Zimmer)and Emerita portoricensis Schmidt. Ecology, 46, 195-197. Gordan, J. (1957).A bibliography of the Order Mysidacea. Bulletin A w & a n Museum of Natural History, 112, 283-393. Gordon, I. (1967).On Spekogriphua, a new cavernicolouscrustacean from South Africa. Bulletin of the British Muaeum (Natural History), Zoology, 5, 31-47.

280

THE BIOLOGY OF MYSIDS

Gordon, I. (1958). A new subterranean crustacean from the West Indies. Nature, London, 181, 1552-1553: Gordon, I. (1960). On a Stygiomyh from the West Indies, with a note on Spelaeogriphus (Crustacea, Peracarida). Bulletin of the British Museum (Natural History),Zoology, 6, 285-324. Gordon, I. (1964). On the mandible of the Stygocaridae (Anaspidacea) and some other Eumalacostraca, with specialreference to the lacinia mobilis. Crustaceana, 7, 150-157. Gourret, P. (1900). Schizopodes. I n “Zoologie Descriptive” (Ed. L. Bouron), 2, pp. 187-213. Grabe, S. A. (1978). Food and feeding habits of juvenile Atlantic tomcod, Microgadus tomcod, from Haverstraw Bay, Hudson River. Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 76, 89-94. Grainger, E. H. (1962). Zooplankton of Foxe Basin in the Canadian Arctic. Journal of the Fisheries Research Board of Canada, 19, 377-400. Grainger, J. N. R. (1956). Effects of changes of temperature on the respiration of certain Crustacea. Nature, London, 178, 930-931. Grant, G. C. (1962). Predation of bluefish on young Atlantic menhaden in Indian River, Delaware. Chesapeake Science, 3, 45-46. Green, J. W. (1970). Observations on the behaviour and larval development of Acanthomysis sculpta (Tattersall) (Mysidacea). Canudian Journal of Zoology, 48, 289-292. Greve, W.(1975). The Meteor planktonkuvette: a device for the maintenance of macrozooplankton aboard ships. Aquaculture, 6, 77-82. Grice, G. D. and Hart, A. D. (1962). The abundance, seasonal occurrence and distribution of the epizooplankton between New York and Bermuda. Ecological Monogruphs, 32, 287-309. Griffiths, W. E. (1976). Food and feeding habits of European perch in the Selwyn River, Canterbury, New Zealand. New Zealand Journal of Marine and Freshwater Research, 10, 417-428. Grimas, U. and Nilsson, N.-A. (1966). On the food chain in some north Swedish river reservoirs. Report of the Institute of Freshwater Reaearch, Drottningholm, 46, 31-48.

Grindley, J. R. and Penrith, M. J. (1965). Notes on the bathypelagic fauna of the seas around South Africa. Zoologica Africana, 1, 275-295. Grossnickle, N.E. and Beeton, A. M. (1979). Antennal scale length as a measure of relative size in the opossum shrimp, Mysis relicta Loven. Crustaceana, 36, 14 1-1 46.

Grossnickle, N. E. and Morgan, M. D. (1979). Density estimates of My& relictu in Lake Michigan. Journal of the Fisheries Research Board of Canada, 36, 694-698.

Gum, G. W. (1978). Antennal sensilla of N e o m y h integer (Leach). Protoplasm, 95, 145-162. Haahtela, I.(1964). Havaintoja Perameren selkarangattomista. Luonnon Tutkija, 68, 162-166.

Haahtela, I. (1973). Litoraalivyohykkeen tutkimisesta Itiimeren talviolosuhteissa. fimnologisymposium, 1970, 73-84. Hacker, V. A. (1956). Biology and management of lake trout in Green Lake, Wisconsin. Transactions American Fisheries Society, 86, 7 1-83. Haedrich, R. L. and Henderson, N. R. (1974). Pelagic food of Coryphaenoides amnatue, a deep benthic rattail. Deep-sea Research, 21, 739-744.

REFERENOES

281

Haefner, P. A. (1968). Occurrence of Mysis gmpensis (0.S. Tattersall) (Mysidacea) in the Gulf of Maine. Crustdceana, 14, 219-220. Haefner, P. A. (1969). Occurrence of a larval alligator fish (Agonidae) in brackish water. Copeia, 201-202. Haertel, L. and Osterberg, C. (1967). Ecology of zooplankton, benthos and fishes in the Columbia River Estuary. Ecology, 48, 459-472. Haidar, Z. (1963). Oecologie du rouget barbet (Mullus barbatus L.) en Adriatique orientale. Rapport et proces-verbaux dea rduniona. Commiesion internatwnale pour l’explorationsoientij2que de la mer Mdditerranhe, 17,363-364. Hair, J. R. (1971). Upper lethal temperature and thermal shock tolerances of the opossum shrimp, Neomysis awatscheds, from the Sacramento-San Joaquin Estuary, California. California Fish and Game, 57, 17-27. Hakala, I. (1978). Distribution, population dynamics and production of Mysis relicta (LovBn) in southern Finland. Annales Zoologici Fennici, 15, 243258.

Hallberg, E. (1977). The fine structure of the compound eyes of mysids (Crustacea: Mysidacea). Cell and Tissue Research, 184, 45-65. Hamond, R. (1974). The marine and brackiRh-water non-amphipodan peracaridan Crustacea of Norfolk. Cahiers de Biologie marine, 15, 197-213. Hand, C. H. and Berner, L. (1960). Food of the Pacific sardine (Sardinops caerulea).Fkhery Bulletin, Fish and Wildlife Service, United States Department of Interior, 60, 175-184. Hanson, J. A. (1966). Final introduction of the opossum shrimp (Mysis relicta Loven) into California and Nevada. California Fish and Game, 52, 220. Hanstrom, B. (1948). The brain, the sense organs, and the incretory organs of the heed in the Crustacea Malacostraca. Bulletin Biologique de la France et de In Belgique, 33, 98-126. Harder, W. (1968). Reactions of plankton organisms to water stratification. fimnology and Oceanography, 13, 156-168. Hardy, A. C. (1924). The herring in relation to its animate environment. Part I. “he food and feeding habits of the herring with special reference to the east coast of England. Fishery Investigations, Ministry of Agriculture, Food and Pisherim, Ser. 11, 7 , 1-53. Hargrave, B. T. and Geen, G. H. (1968). Phosphorus excretion by zooplankton. fimnology and Oceanography, 13, 332-342. Hartley, C. H. and Fisher, J. (1936). The marine foods of birds in an inland fjord region in western Spitsbergen. Part 2. Journal of Animal Ecology, 5, 370-389.

Hartley, P. H. T. (1940). The Saltash tuck-net fishery and the ecology of some estuarine fishes. Journal of the Marine Biological Association of the United Kingdom, 24, 1-68. Hartnoll, R. G. (1978). The effect of salinity and temperature on the post-larval growth of the crab Rhithropanopeus harrisii. I n “Physiology and Behaviour of Marine Organisms” (Eds D. S. McLusky and A. J. Berry), pp. 349-358. Pergamon Press, Oxford and New York. Harvey, E. N. (1952). “Bioluminescence”, 632 pp. Academic Press, New York and London. Hatzakis, A. (1977). Contribution a 1’6tude des Gastrosaccinae (Crustacea, Mysidacea) de la MBditerranee. Description de Haplostylus bucescui n.sp. et revision de la nomenclature des Haplostylus et Qagtrosaccus MBditerran6ens. Biologia CfaWo-Helleniea,6, 271-287.

282

THE BIOLOQY OF MYSIDS

Hatzakis, N. (1974). Mysidac6s trouv6s en Mer Eg6e (partie sud) de 1966 h 1971. Rapport et proces-verbaux des rkunions. Commiasion internationale pour l’exploratim scienti$que de la mer Mkditerranke, 22,81. Havinga, B. (1941). De veranderingen in den hydrographischen toestand en in de macrofauna van het Ijsselmeer gedurende de jaren 1936-1940. De Biologie van de Zuiderzee Tijdens haar Drooglegging, 5 , 5-26. Healey, M. C. and Woodall, W. L. (1973). Limnologcal surveys of seven lakes near Yellowknife, Northwest Territories. Fisheries Research Board of Canada, Technical Report 407, 1-34. Henderson, J. R. (1885). Recent additions to the invertebrate fauna of the Firth of Forth. Proceedings of the Royal Phykcal Society of Edinburgh, 8, 307-315. Henderson, J. T. (1927). A note on the effect of temperature on the cardiac rhythm of certain schizopods. Journal of experimental Biology, 5 , 135-137. Herdman, W. A. (1890). Third Annual Report of the Liverpool Marine Biological Station on Puffin Island. Proceedings and Transactiona of the Liverpool Biological Society, 4, 3 6 7 4 . Herdamn, W. A. (1920).List of species recorded by the Liverpool Marine Biological Laboratory since the publication of the British Association list in 1896. Proceedings and Transactions of the Liverpool Biological Society, 34, 75-78. Herdman, W. A., Thompson, I. C. and Scott, A. (1898). On the plankton collected continuously during two traverses of the North Atlantic in the summer of 1897; with descriptions of new species of Copepoda; and an Appendix on dredging in Puget Sound. Proceedings and Transactions of the Liverpool Biological Society, 12, 33-90. Herman, S. S. (1962). Spectral sensitivity and phototaxis in the opossum shrimp, Neomysis americana Smith. Biological Bulletin, Marine Biological Laboratory, Wood’s Hole, Mass. 123, 562-570. Herman, S. S. (1963). Vertical migration of the opossum shrimp Neomysis americana Smith. Limnology and Oceanography, 8, 228-238. Herman, S. S., Mihursky, J. A. and McErlean, A. J. (1968). Zooplankton and environmental characteristics of the Patuxent River Estuary, 1963-1965. Chesapeake Science, 9, 67-82. Herring, P. J. (1974). Size, density and lipid content of some decapod eggs. Deep-Sea Research, 21, 91-94. Hesthagen, I. H. (1973). Diurnal and seasonal variations in the near-bottom fauna-the hyperbenthos-in one of the deeper channels of the Kieler Bucht (western Baltic). Kieler Meeresforschungen, 29, 116-140. Hesthagen, I. H. and Gjermundsen, B. (1978). The replicability of sampling the hyperbenthic region by means of Beyer’s 50cm epibenthic closing net. Meeresforschung, Reports on Marine Research, 26, 1-10. Hesthagen, I. H. and Gjermundsen, B. (1979). Late summer dipma1 migration in the hyperbenthos of Vejsnas Rinne, Western Baltic. Meeresforachung, Reports on Marine Research, 27, 19-26. Heubach, W. (1969). Neomysk awatschem-s in the Sacramento-San Joaquin River Estuary. Limnology and Oceanography, 14, 533-546. Heubach, W., Toth, R. J. and McCready, A. M. (1963). Food of young-of-the-year striped bass (Roccus saxatilis) in the Sacramento-San Josquin River System. California Fish and Game, 49, 224-239. Hewitt, C. G. (1907). Ligia. Liverpool Marine Biology Committee, Memoir, 1 4 , 1 4 7 . Hiatt, R. W. and Strasburg, D. W. (1960). Ecological relationships of the fish fauna on coral reefs of the Marshall Islands. Ecological Monographs, 3 0 , 6 6 1 2 7 .

REFERENCES

283

Hirrtno, T. (1975). On the distribution and seasonal variation of homarine in some marine invertebrates: Bulletin of the Japanese Society of Scienti$c Fisheries, 41, 1047-1051.

Hirano, T., Kikuchi, T. and Okada, I. (1964a). Contents of inorganic substance and vitamin B,, in Euphausia. Bulletin of the Japanese Society of Scientific Fisheries, 30, 267-271. Hirano, T., Kikuchi, T., Ino, K., Tanoue, N., Taguchi, T. and Okada. I. (1964b). Contents of inorganic substance and vitamin B,, in Euphamia. Journal of the Tokyo University of Fisheries, 50, 65-70. Hjort, J. and Dahl, K. (1900). Fishing experiments in Norwegian fjords. Report on Norwegian Fishery and Marine Investigations, 1, 1-215. Hobson, E. S. (1974). Feeding relationships of teleostean fishes on coral reefs in Kona, Hawaii. Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 72, 915-1031. Hobson, E. S. and Chess, J. R. (1976). Trophic interactions among fishes and zooplankters near shore at Santa Catalina Island, California. Fishery Bulktin, National Oceanic Atmospheric Administration of the United States, 74, 567-598. Hobson, E. S. and Chess, J. R. (1978). Trophic relationships among fishes and plankton in the lagoon of Enewetak Atoll, Marshall Islands. Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 76, 133-153. Hodge, D. (1963a). A new species of mysid (Mysidacea: Crustacea) from the Brisbane River. University of Queensland Papers, Department of Zoology, 11, 81-88.

Hodge, D. (1963b). The distribution and ecology of mysids in the Brisbane River. University of Queensland Papers, Department of Zoology, 11, 89-104. Hodge, D. (1964). A redescription of Tenagmysia chiltoni (Crustacea: Mysidacea) from a freshwater coastal lake in New Zealand. New Zealand J o u d of Science, 7, 387-395. Hoenigman, J. (1954). Novosti s Podrocja Jadranskega Zooplanktona. Nouveaut6s concernantes le zooplankton Adriatique (sur la trouvaille des Ellobiopside). Bioloski Vestnik, Ljubljana, 3, 1 0 6 116. Hoenigman, J. (1958a). Quelques observations sur les “zoocourants” dans I‘Adriatique. Rapport et prom-verbaux des rbunwna. Commission internationale pour l’exploration scientiijique de la mer Mkditewanke, 14,261-262. Hoenigman, J . (195813). Sur la dbcouverte de quelques especes zooplanktoniques nouvelle pour I’Adriatique, de deux Bpibiontes nouveaux pour les mysidads, et de deux especes de parasites nouvelles pour le domaine MBditerranBen. Rapport et proces-verbaux dea rkunions. Commieaion interntationale pour I’exploration scientifiue de la mer Mdditerrande, 14, 263-264. Hoenigman, J. (1960). Faits nouveaux concernant les mysidaces (Crustacea) et leurs epibiontes dans YAdriatique. Rapport et procee-verbaux dea r k u n i m . Commi.ssion internationale pour l’exploration scientihue de la m r Mdditerrande, 15, 339-343.

Hoenigman, J. (1961). Sur quelques esphces nouvelles pour YAdriatique. Rapport et proces-verbaux des r k u n d . Commission intemtcttionale pour ,?’exploration scientifque de la mer Mdditerrande, 16, 217-218. Hoenigman, J. (1963). Mysidacea de 1’Expedition “Hvar” (1948-49) dans 1’Adriatique. Rapport et proces-verbaux dee rbunwna. Commission internationale pour l’exploration scientihue de la mer Mdditerrande, 17, 603-616. Hoenigman, J. (1964a). Baetrosaccwr sanctm (Van. Ben.) et sa relation possible avec la genese de 1’Adriatique. Acta Adriatica, 11, 135-143.

284

THE BIOLOGY OF MYSIDS

Hoenigman, J. (1964b). Sur quelques facteurs importants de la repartition horizontale du zooplankton Adriatique. Acta Adriatica, 11, 145-160. Hoenigman, J. (1965). Note sur les copepodes, leurs epibiontes (Suctoridae) et sur un Mysidach, h8te nouveau de 1’Amallocystis (Ellobiopsidae). Rapport et proces-verbaux des rdunions. Cornmiasion internatwnale pour l’exploration scientifique de la m r Mdditerrande, 18, 473. Hoenigman, J. (1968). Sur les mysidac6s des eaux ouvertcs de 1’Adriatique septentrionale. Rapport et proem-verbaux des riunions. Commission internationale pour l’exploration scientiflque de la mer Mdditemane’e, 19, 449. Hoffman, E. G. and Yancey, R . M. (1966). Ellobiopsidae of Alaskan coastal waters. Pacific Science, 20, 70-78. Hogstad, A. M. (1969). The X-organ and the sinus gland of Mysis relicta Loven (Crustacea Mysidacea). Nytt Magasin for Zoologi, 17, 105-109. Holland, A. F. and Polgar, T. T. (1976). Seasonal changes in the structure of an intertidal community. Marine Biology, 37, 341-348. Holmquist, C. (1949). Uber eventuelle intermediare Formen zwischen Mysis oculate Fabr. und Mysis relicta Loven. Lunds Universitets Arsskrift, Ser. 2, 45, 1-25. Holmquist, C. (1955). Die “Mysis relicta” aus dem Mittelmiergebiet ( = Dianzysis bahirensis (G. 0. Sam) 1877 und Puramysis helleri (G. 0. Sam) 1877). Zoologischer Anzeiger, 154, 277-288. Holmquist, C. (1956). Betrachtungen uber Boreomysis rostrata Illig und weitere Boreomysis-Arten. Arkiv f or Zoologi, Ser. 2, 10, 427-447. Holmquist, C. (1957a). On aberrant specimens of Praunus Jexuosus and some other opossum shrimps. Acta Borealis, 13, 1-29. Holmquist, C. (195713).Mysidacea of Chile. Reports of the Lund University Chile Expedition 1948-1949 (Report 28). Lunds Universitets Arsskrift, Ser. 2, 53, 1-52. Holmquist., C. (195th). On a new species of the genus Mysis, with some notes on Mysw oculatu (0.Fabricius). Meddelelser om Gronland, 159 (4), 1-17. Holmquist, C. (195%). Proposed use of the plenary powers to validate a neotype for the nominal species “Cancer oculatus” Fabricius (O.), 1780, to designate the species so named to be the type species of the genus “Mysis” Latreille, 1802-1803 (Class Crustacea, Order Mysidacea) and matter incidental thereto. Bulletin of Zoological Nomenclature, 16, 51-61. Holmquist, C. (1959a).Mysis polaris-a new species of the genus Mysis. Kungliga E’ysiograJiska sallskapets i Lund Forhandlingar, 29, 21-25. Holmquist, C. (1959b). “Problems on marine-glacial relicts on account of investigations on the genus Mysis”, 270 pp. Berlingska Boktryckeriet, Lund. Holmquist, C. (1962). The relict concept--is it merely a zoogeographic conception. Oikos, 13, 262-292. Holmquist, C. (1963a). Some notes on Mysis relicta and its relatives in northern Alaska. Arctic, 16, 109-128. Holmquist, C. (1963b). Dags a t t omvardera de s. k. marin-glaciala relikterna. Fauna och Flora, 30-42. Holmquist, C. fl966). Die sogenannten marin-glazialen. Relikte nach neueren Gesichtsponkten. Archiv f fir Hydrobiologie, 62,285-326. Holmquist, C. (1973). Taxonomy, distribution and ecology of the three species Neomysis intermedia (Czerniavsky), N . awatschensis (Brandt) and N . mercedis Holmes (Crustacea, Mysidacea). Zoologische Jahrbucher, S y s t e w t i k , OkOlOgie und Geographie der Tiere, 100, 197-222.

REFERENCES

285

Holmquist, C. (1975).Arevision of the speciesArchaeomysi8grebnitzkii Czerniavsky and A . maculata (Holmes) (Crustacea, Mysidacea). Zoologische Jahrbucher, Systematik, Okologie und Beographie der Tiere, 102, 51-71. Holt, E. W. L. and Beaumont, W. I. (1899). On Siriella armata (M.-Edw.) and the reputed occurrence of 8. frontalis (M.-Edw.) in British seas. Annaba and Magazine of Natural History, Ser. 7, 3, 151-152. Holthuis, L. B. (1967). Mysidacea, Cirripedia and Cladocera newsletters. Crmstmeanu, 12, 110-1 11. Holthuis, L. B. and Sivertsen, E. (1967). The Crustacea Decapoda, Mysidacea and Cirripedia of the Tristan da Cunha Archipelago with a revision of the “frontaZis” subgroup of the genus J w . Results of the Nomuegian Scienti$c Expedition to Trktan da Cunha, 1937-1938, 52, 1-55. Hopkins, C. C. E. and Gulliksen, B. (1978). Diurnal vertical migration and zooplankton-epibenthos relationships in a north Norwegian fjord. I n “Physiology and Behaviour of Marine Organisms” (Eds D. S. McLusky and A. J. Berry), pp. 271-280. Pergamon Press, Oxford. Hopkins, T. L. (1958). Mysid shrimp. Estuarine Bulletin, University of Delaware, 3, 4-6. Hopkins, T. L. (1965). Mysid shrimp abundance in surface waters of Indian River Inlet, Delaware. Chesapeake Science, 6, 86-91. Hopkins, T. L. (1966). The plankton of the St Andrew Bay System, Florida. Publicatiow of the Institute of Marine Science, Univer&y of Texm, 11, 12-64. Hulburt, E.M. (1957). The distribution of Neomysis americana in the estuary of the Delaware River. Limnology and Oceanography, 2, 1-11. Hureau, J. C. and Monod, T. (1973). Check list of the fishes of the north-eastern Atlantic and of the Mediterranean, 1, 683 pp. United Nations Educational, Scientific and Cultural Organization, Paris. Hynes, H. B. N., Macan, T. T. and Williams, W. D. (1960). A key to the British species of Crustacea : Malacostraca occurring in fresh water. Scienti$c Publicatiom of th,e Freshwater Biological Association, 19, 1-36. Ii, N. (1964a). “Fauna Japonica, Mysidae (Crustacea)”, 610 pp. Biogeographical Society of Japan. Ii, N. (1964b). Report on a small collection of Mysidacea from coastal waters of Asamushi. Bulletin of the Biological Station of Asamuahi, Tohoku University, 12, 1-7. Ikeda, T. (1970). Relationship between respiration rate and body size in marine plankton animals as a function of the tempereture of habitat. Bulletin of the Faculty of Fisheries, Hokkaido University, 21, 91-112. Ikeda, T. (1977). The effect of laboratory conditions on the extrapolation of experimental measurements to the ecology of marine zooplankton. 11.Effect of oxygen saturation on the respiration rate. Bulletin of the Plankton Society of Japan, 24, 19-28. Illig, G. (1905). Das Leuchten der Gnathophausien. Zoologischer Anzeiger, 28, 662. Ingle, R . W. (1972). A redescription of Spelaeomysis servatua (Fage) comb. nov. (Mysidacea: Lepidomysidae) from the material collected on Aldabra Atoll, with a key to the species of Lepidomysidae. Bulletin of the British Muaeum (Natural History), Zoology, 22, 199-210. hyang, N. M. (1978). Notes on food of the Baltic palaemonid shrimp, Palaemon adsperm var. fabricii (Rathke). Meeresforschung,Reports on Marine Research, 26, 42-46.

286

THE BIOLOGY OF MYSIDS

Isaacs, J. D., Fleminger, A. and Miller, J. K. (1969). Distributional atlas of zooplankton biomass in the California Current region: spring and fall 19551959. California Cooperative Oceanic Fisheries Investigations, Atlas 10, 1-252. Ishikawa, M. and Oshima. Y . (1951). On the life history of a mysidcrustacean, Neomysis Japonica Nakazawa. Bulletin of the Japanese Society of Scientific Fisheries, 16, 461-477. Ivanov, S. N. (1972). Introduction of mysids into ponds as a means of increasing fish productivity. Hydrobiology Journal, 8 , 24-30. Jacobs, F. and Grant, G. C. (1974). Acute toxicity of unbleached kraft mill effluent (UKME) to the opossum shrimp, Neomy& americana Smith. Water Research, 8, 439-445. Jander, R. (1962). The swimming plane of the crustacean Mysidium gracile. Biological Bulletin, Marine Biological Laboratory, Wood’a Hole, Mass., 122, 380-390.

Jander, R. and Waterman, T. H. (1960). Sensory discrimination between polarized light and light intensity patterns by arthropods. Journal of Cellular and Comparative Physiology, 56, 137-159. Jansson, B. 0. and Kiillander, C. (1968). On the diurnal activity of some littoral peracarid crustaceans in the Baltic Sea. Journal of Experimental Marine Biology and Ecology, 2, 24-36. Jawed, M. (1969). Body nitrogen and nitrogenous excretion in Neomysis rayii Murdoch and Euphausia paci$ca Hansen. Limnology and Oceanography, 14, 748-754.

Jawed, M. (1973). Effects of environmental factors and body size on rate9 of oxygen consumption in Archaeomysis grebnitzkii and Neomysia awatscheneis (Crustacea: Mysidae). Marine Biology, 21, 173-179. Jensen, J. P. (1958). The relation between body size and number of eggs in marine malacostraces. Medddelser fra Danmarks Fiakeri+g Havundersogeleer, 2 (19), 1-25.

Jepsen, J. (1965). Marsupial development of Boreomysis arctica (Kroyer, 1861). Saraia, 20, 1-8. Jespersen, P. (1928). Investigations on the food of the herring in Danish waters. Meddelelser fra Kommissionen for Havunderssgelaer, Seriee Plankton, 2 (2), 1-149.

Jespersen, P. (1932). On the food of the herring in Icelandic waters. Meddeklaer fra Kommissionenfor Danmarks Fiskeri-og Havunders0gelser, Series Plankton, 2 (3), 1-34.

Jespersen, P. (1936). The food of the herring in the waters round Bornholm. Meddelelser fra Kommissionen for Dunmark8 Fiskeri-og Havunders0gelaer, Serie Plankton, 3 (2), 1-40. Johnson, C. R. (1969). Contributions to the biology of the showy snailfish, Liparb pulchellus (Liparidae). Copeia, 830-835. Johnson, J. T. and Hopkins, T. L. (1978). Biochemical components of the mysid shrimp Taphromysis bowmani Bacescu. Journal of Experimental Marine Biology and Ecology, 31, 1-9. Johnson, L. (1964). Marine-glacialrelicts of the Canadian Arctic islands. Systematic Zoology, 13, 76-91. Johnson, L. (1975). Distribution of fish species in Great Bear Lake, Northwest Territories, with reference to zooplankton, benthic invertebrates, and environmental conditions. Journal of the Fiaheries Research Board of Canada, 32, 1989-2004.

287

REFERENUES

Jolicoeur, P. and Brunel, P. (1966). Application du diagramme hexagonal B 1’6tude de la selection de ses proies par la morue. Vie et Milieu, 17, 4 19-433.

Jones, R. (1954). The food of the whiting, and a comparison with that of the haddock. Marine Research, 2, 1-34. Juchault, P. (1963). Sur la glande androghne d‘un certain nombre de PBracarides (cumac6s, mysidacbs, tanaidac6s). Compte Rendu des S h m de la Biologie, 157, 613-615.

Judkins, D. C. and Wright, R. (1974). New records of the mysida Boreomysb nobilis G. 0. Sars and My& litoralis (Banner) in the Saguenay Fjord (St. Lawrence Estuary). Canadian Journal of Zoology, 52, 1087-1090. June, F. C. and Carlson, F. T. (1971). Food of young Atlantic menhaden, Brevoortia tyrannzcs, in relation to metamorphosis. Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 68, 493-512. Just, J. (1970). Decapoda, Mysidacea, Isopoda and Tanaidacea from Jergen Brenlund Fjord, North Greenland. Me&klaer om Q r d a n d , 184 (9), 1-32.

Kaliyamurthy, M. (1972). Notes on the brackish water mysid Mesopodopeis orientalis (Tattersall) of the Pulicat Lake, Journal of the Inland Fkheries Society of India, Barrackpore. 4, 198-200. Kane, J. E. (1964). Thalassomycesnzaraupii, a new species of ellobiopsid parasite on the hyperiid amphipod Parathemieto gaudichudii (Guer.) New Zealand Journal of Science, 7, 289-303. Krtne, W. F. de V. (1904). Further captures of Mysia relicta in Ireland. Irish Naturalist, 13, 107-109. Kasaoka, I. D. (1974). The male genital system in two species of mysid Crustacea. Journal of Morphology, 148, 259-284. Keeble, M. A. and Gamble, F. W. (1903). The colour-physiology of higher Crustacea. Proceedings of the Royal Society of London, 71, 69-71. Keeble, M. A. and .Gamble, F. W. (1904). The colour-physiology of higher Crustacea. Philosophical Tramactions of the Royal Society of London, B, 196,

.

295-388.

Khlebovich, V. V., Bogdanov, V. E. and Kuzmina, V. G. (1970). Euryhalinity and osmoregulation in Mysis oculata and M. littoralis from the White Sea. Zoologischeskii Zhurnal, 49, 1085-1088. Kikuchi, T. (1967). An ecological study on animal communities of the Zostera marina belt in Tomioka Bay, Amakusa, Kyushu. Publications of the Amakwra Marine Biological Laboratory, 1, 1966, 1-106. Kinahan, J. P. (1859). Notes on dredging in Belfast Bay with a list of species. Natural History Review, 6, 79-86. Kinne, 0. (1955). Neomyk vulgaria Thompson eine autokologisch-biologische studie. Biologisches Zentralblatt, 74, 160-202. Kinney, W. L. (1972). The macrobenthos of Lake Ontario. Proceedings of Fijtmnth Conference on Great Lakes Research, 53-79. Kiortsis, V. (1974). Quelques consid6rations sur 1’6cologie du zooplankton en Mer Eg6e. Repartition geographique et variations saisonnidres. Rapport et proces-verbaux des rduniona. Commission intemtwnale pour l’exploration scientifque de la mRr Mdditerranh, 22, 139-141. Kisldioglu, M. and Gibson, R. N. (1976a). Prey “handling time” and its importance in food selection by the 15-spined stickleback, Spinachia spinachia (L.). Journal of Experimental Marine Biology and Ecology, 25, 151-158.

288

THE BIOLOGY OF MYSIDS

Kislalioglu, M. and Gibson, R. (1976b). Some factors governing prey selection by the 15-spined stickleback, Spinachia spinachia (L.).Journal of Experimental Marine Biology and Ecology, 25, 159-169. Klawe, W. L. (1955). Pseudomma a@ne G. 0. Sam: an addition to the list of Mysidacea of Eastern Canada. Canadian Field Naturalist, 69, 66. Klyashtorin, L. B. and Kuz’micheva, V. I. (1975). Level of energy expenditure of planktonic crustaceans on active movements. Oceanology, 15, 592-595. Klyashtorin, L. B. and Musayeva, E. I. (1977). The use of oxygen electrodes for measuring the respiration rate of planktonic animals. Oceanology, 17,595-597. Klyashtorin, L. B. and Yarzhombek, A. A. (1973). On energy losses for active movement of planktonic organisms. Okeanologiya, 13, 697-703. Kohout, F. A. and Kolipinski, M. C. (1967). Biological zonation related to groundwater discharge along the shore of Biscayne Bay, Miami, Florida. I n “Estuaries” (Ed. G. H. Lauff), pp. 488-499. American Association for the Advancement of Science, Washington, D.C. Publication No. 83 Koller, G. and Mayer, E. (1930). Versuche uber den Wirkungsbereich von Farbwechselhormonen. Biologisches Zentralblatt, 50, 759-768. Kost, A. L. B. and Knight, A. W. (1975). The food of Neomysis mercedis Holmes in the Sacramento-San Joaquin Estuary. California Fish and Game, 61,35-46. Kravitz, M. J., Pearcy, W. G. and Guin, M. P. (1976). Food of five species of cooccurring flat fishes on Oregon’s continental shelf. Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 74, 9 8 6 9 9 0 . Kropp, B. and Perkins, E. B. (1933).The occurrence of the humoral chromatophore activator among marine Crustaceans. Biological Bulletin, Marine Biological Laboratory, Wood’s Hole, Mass., 64, 28-32. Krylova, A. G., Romanycheva, 0. D. and Sergieva, Z. M. (1975). Role of zooplankton in the feeding of fish reared in cages in the Rozhok Bight. Trwlg Vsesoyuznogo Nauchnoksledovatd’skogo Instituta Morskogo Rgbnogo Kkazyaistva i Okeanografii, 59, 140-151. Kuhl, H. and Mann, H. (1972). Untersuchungen ZUT Hydrobiologie der unteren Ems. Archiv f ilr Fischereiwksenschaft, 23,243-268. Kuhlmorgen-Hille, von G. (1963). Quantitative untersuchungen der Bodenfauna in der Kieler Bucht und ihre jahreszeitlichen Veriinderungen. Kiekr Meeresforschungen, 19, 42-66. Kulakovskii, E. E. (1969). Neurosecretory system of Mysis oculata. DokZudg A k d m i i nauk SSSR, 192, 226-228. Kulakovskii, E. E. (1971). Neurohormonal control for chromatophores in mysids. Dokladg Akademii nauk SSSR, 196, 234-236. Kulakovskii, E. E. (1974). Neurosecretory system of mass species of mysids from the White Sea. I n “Explorations of the Fauna of the Seas. XI11 (XXI). Seasonal phenomena in the life of the White and Barents Seas”, pp. 290-306. Zoological Institute of the Academy of Sciences of the USSR. Kulakovskii, E. E. (1978). Some basic regularities in the development of the neurosecretory system in crustaceans. Zhumal obshchi biologii, 39, 297-303. Labat, R. (1954). Observations sur l’accouplement et la ponte de Paramy& nouveli. Bulletin de la Socidtd d’HLvtoire Naturelle de Todouae, 89, 406-409. Labat, R. (1957). Observations sur le cycle annuel et la sexualit6 de Paramysis nouveli et Paramysie baceacoi (CrustacBs Mysidac6s) dens la region de Roscoff. Archives de Zoobgie Exprimentale et Gdndrale, 94, 162-173. Labat, R. (1959). Sur le nombre chromosomique chez trois esphces de Mysidach. Bulktin de la Socidtd d’Histoire Naturelle de Toulouae, 94, 201-202.

REFERENCES

289

Labat, R. (1961). L’appareil genital male de Praunus jlexuosw, (Crustace, Mysidace). Bulletin de la Socidtd d’Histoire Naturelle de Toulouse, 96, 61-66.

Labat, R. (1962). La spermatogenhse chez Praunusjlexuusw, (Crustace, MysidacB). Bulletin de la Socidtd d’Histoire Naturelle de Toulouae, 97, 51-60. Lagardhre, J.-P. (1972). Recherches sur l’alimentation des crevettes de la pente continentale Marocaine. Tethys, 8, 655-675. LagardBre, J.-P. (1973). Donn6es sur la biologie et s u r l’alimentation de Dichelopandalw, bonnieri (CrustacB-Natantia) dam le Golfe de Gascogne. Tethys, 5 , 155-166. LagardBre, J.-P. (1977a). Recherches Bur le regime alimentaire et le comportement pddateur des d6capodes benthiques de la pente continentale de 1’Atlantique nord oriental (Golfe de Gascogne et Maroc).I n “Biology of Benthic Organisms” (Eds B. F. Keegan, P. 0. Ceidigh and P. J. S. Boaden), pp. 397-408. Pergamon Press, Oxford. Lagardh, J.-P. (1977b). Recherches sur la distribution verticale et sur l’alimentation des crustac6s decapodes benthi,ques de Is, pente continentale du Golfe de Gascogne. Analyse des groupements carcinologiques. Rullet
290

THE BIOLOGY OF MYSIDS

Lebour, M. V. (1923). The food of plankton organisms 11.Journal of the Marine Biological Association of the United Kingdom, 13, 70-92. Ledoyer, M. (1963). Hemimysis speluncola n.sp. mysidace nouvelle des grottes sous marines obscures. Recueil des Travaux de la Station Marine d’Endoum, Facultk des Sciences de Marseille, 30, 77-81. Ledoyer, M. (1965). Note sur la faune vagile des grottes sous-marines obscurefi. Rapport et proces-verbaux des re’unions. Commission internationale pour l’exploration scientifique de la mer Mditerranke, 18, 121-124. Ledoyer, M. (1966). Ecologie de la fauns vagile des biotopes MBditerranBens accessibles en scaphandre autonome. I. Introduction donnbes analytiques sur sur les biotopes de substrat dur. Recueil des Travaux.de la Station Marine d’Endoum, Facultk des Sciences de Marseille, 40, 103-149. Ledoyer, M. (1969). Sur divers crustaces Antarctiques (LeptostracBs, CumacBs, MysidacBs et Carid&) recueillis en Terre Adelie en 1961-1963 et 1964-1965. Crustaceana, 17, 88-96. Ledoyer, M. (1970). Mysidacbs des herbiers de phanerogames marines de Tulear (Madagascar) Btude systematique et Bcologique. Recueil des Tram= de la Station Marine d’Endoum, Facultd des Sciences de Marseille, Supplement 10, 2 2 3-2 2 7. Ledoyer, M. (1974). Anisomysis vameuri n.sp. MysidacB nouveau vivant it l’entr6e les grottes sous-marines recifales. Tethys, 5, 1973, 361-366. Ledoyer, M. (1977). Ceratonzyek ericula n.sp. (Crustacea, Mysidacea) recolt6e au large des Iles Kerguelen. Bulletin d u Museum National d’Histoire Naturelle, Series 3, Zoologic, No. 432, 253-258. Lee, B. D. and Chin, P. (1971). Studies on the biochemical composition of the mysid, Neomysis awatschensis (Brandt ), Publications of the Marine Laboratory, Pusan Fisheries College, 4, 1-8. Lee, C. Y. and Chin, P. (1976). Effects of temperature and starvation on the oxygen consumption and nitrogen excretion of a mysid, Paracanthomysis hispida Ii. Publications of the Institute of Marine Science, National Fkhem’es University of Buaan, Korea, 9, 25-31. Lee, R. F. and Hirota, J. (1973). Wax esters in tropical zooplankton and nekton and the geographical distribution of wax esters in marine copepods. Lintnology and Oceanography, 18,227-239. Lee, R.F., Hirota, J. and Barnett, A. M. (1971). Distribution and importance of wax esters in marine copepods and other zooplankton. Deep-sea Research, 18, 1147-1 165. Leslie, G. and Herdman, W. A. (1881). The invertebrate fauna of the Firth of Forth, Proceedings of the Royal Physical Society of Edinburgh, 6, 201-231. Le Sueur, R. F. (1954). The Cumacea, Mysidacea, Amphipoda, Isopoda and Tanaidacea of the Channel Islands. Bulletin Annuel de la Socidtd Jer&zke, 16, 207-216. Lettevall, A. U. (1962). Mysis relicta, Pontoporeia a f i n k , Pallasea quadrispinoea och Mesidotea entomon funna i. sjoar i mellersta Kalmar liin. Fauna och Flora, Uppsala, 66-76. Lewis, R. W. (1967). Fatty acid composition of some marine animals from various depths. Journal of the Fisheries Research Board of Canada, 24, 11011115. Liao, H.-H. (1951). Seasonal tidal and diurnal movements and breeding of the free-swimming Crustacea Malacostraca of Port Erin Bay. Thesis for the degree of Ph.D., University of Liverpool.

REFERENCES

291

Light, S. F. (1954). Intertidal Invertebratm of the Central Californicc Coast”, 446 pp. University of California Pres8, Berkeley. Lindquist, A. (1959). Studien uber das zooplankton der Bottensee 11. Zur verbreitung und Zusammensetzung des zooplanktons. Report of the Institute of Marine Research, Lysekil, Series Biology, No. 11, 1-136. Linford, E. (1965). Biochemical studies on marine zooplankton 11. Variations in the lipid content of some Mysidacea. Journal du Conseil, 30, 16-27. Linn, J. D. and Frantz, T. C. (1965). Introduction of the opossum shrimp (My& relktu Loven) into California and Nevada. Cdifornicc P k h and Game, 51,4&51. Lochhead, J. H. (1950). Heteromysis fomnosa. I n “Selected Invertebrate Types” (Ed. F. A. Brown), pp. 418-422. John Wiley and Sons, Inc., New York. Longhurst, A. R. (1957). The food of the demersal fkh of a West African estuary. Journal of Animal Ecology, 26, 369-387. LBnnberg, E. (19314. Untersuchungen uber das Vorkommen carotinoider Stoffe bei marinen Evertebraten. Arkiv fiir Zoologi, 2ZA (la), 1-49. Lijnnberg, E. (1931b). Zur Kenntnis der Carotinoide bei marinen Evertebraten. Arkiv for Zoologi, 23A (15), 1-74. Lamberg, E. (1934). Witere Beitriige zur Kenntnis der Carotinoide der marinen Evertebraten. Arkiv for Zoobgi, 26 (7), 1-36. Love, Y.S. and Ebeling, A. W. (1978). Food and habitat of three switch-feeding fishes in the kelp forests off Santa Barbara, California. F k h r y Bulletin, National Oceanic Atmosphric Administration of the United States, 76, 257-27 1. Lucas, 6. E. (1936). On certain inter-relations between phytoplankton and zooplankton under experimental conditions. Journal du Conseil, 11, 343-362. Lucu, C. (1978). Sodium balance and salinity tolerance of the mysid L e p t o m y k mediterranea. I n “Physiology and Behaviour of Marine Organisms” (Eds D. S. McLusky and A. J. Berry), pp. 95-103. Pergamon Press, Oxford. Macdonald, A. G. and Gilchrist, I. (1978). Further studies on the pressure tolerance of deep-sea Crustacea, with observations using a new high pressure trap. Marine Biology, 45, 9-21. McEachran, J. D., Boesch, D. F. and Musick, J. A. (1976). Food division within two sympatric species-pairs of skates (Pisces: Rajidae). Marine Biology, 35, 301-31 7. Macer. C. T. (1967). A new bottom-plankton sampler. Journal du Coneeil, 31, 168-163. McHugh, J. L. (1967). Estuarine nekton. I n “Estuaries” (Ed. G . H. Lauff), pp. 581-620. American Association for the Advancement of Science, Washington, D.C., Publication No. 83. McIntosh, W. C. (1919). Notes on some of the Peracarida (Mysidacea) and Eucarida (Decapoda)of St Andrews, in the living condition. Scottish Naturalist, 79-86. Mackintosh, N. A. (1934). Distribution of the macroplankton in the Atlantic sector of the Antarctic. “Discovery” Reports, 9, 65-160. McLachlan, A., Wooldridge, T. and Horst, G. van der (1979). Tidal movements of the macrofauna on an exposed sandy beach in South Africa. Journal of Zoobgy, 187, 433-442. McLaren, I. A. (1958). The biology of the ringed seal (Phoca hisppida Schreber) in the eastern Canadian Arctic. Bulletin of the Fisheries Research Board of Canada, 118, 1-97. McLellan, T. (1977). Feeding strategies of the macrourids. Deep-sea Research, 24, 1019-1036.

292

THE BIOLOGY OF MYSIDS

McLusky, D. S. and Heard, V. E. J. (1971). Some effects of salinity on the mysid Praunua jlexuosus. Jo&nal‘of the Marine Biological Association of the United Kingdom, 51, 709-715. McNaught, D. C. (1968). Acoustical determination of zooplankton distributions. Proceedings of the Eleventh conference on Great Lakes Research, 76-84. McNaught, D. C. and Hasler, A. D. (1966). Photoenvironments of planktonic Crustacea in Lake Michigan. Verhandlungen der Internationalen Vereinigung f u r theoretiache und angewandte Limnologie, 16, 194-203. Macpherson, E. (197th).RBgimen alimentario de Micromeeietiua poutmsou (Risso, 1810) y Gadiculua argenteua argenteua Guichenot, 1850 (Pisces, Gadidae) en le Mediterrheo occidental. Inveatigacion P q u e r a , 42, 305-3 16. Macpherson, E. (1978b). RBgimen alimentario de Syvmphurwr nigrescens (Pisces, Cynoglossidae) en el Mediterrheo occidental. Investigacion Peaquera, 42, 325-333. Macpherson, E. (1978~).RBgimen alimentario de Phycis blennoides (Briinich) y Antonogadus megalokynodon (Kolombatovic) (Pisces: Gadidae) en el Mediterr h e o occidental. Investigacion Pesquera, 42, 455-466. Macquart-Moulin, C. (1965). Les mysidacBs bentho-planktoniques du Golfe de Marseille. Recueille des Travaux 0% la Station Marine d’Endoume, Facultk des Sciences de Marseille, 38, 129-253. Macquart-Moulin, C. (1970). Le comportement d’essaim chez les mysidacBsobservation et analyse de quelques essaima littoraux. Rapport et pooeclverbaux de rdunions. Commiesion internationale pour 1’exploration acientifiue de la mer M6ditewandee,20,439-441. Macquart-Moulin, C. (1972). Modifications des rBactions photocinBtiques des Peracarides de l’hyponeuston nocturne en fonction de l’importance de I’eclairement. Tetlrys, a, 1971, 897-920. Macquart-Moulin, C. (1973a). Le compoeement d’essaim chez les mysidacBs: influence de l’intensitk lumineuse s u p la formation, le maintien e t la dissociation des essaims de Leptomysis lingwra. Rapport et proces-verbaux dea rdunions. Commission intermtionale pour l’explwation acientifiue de la naer Mkditerrande, 499-501. Macquart-Moulin, C. (l973b).L’activitB natatoire rhythmique chez les peracarides bentho-planktoniques. Determinisme endoghe des rhythmes nycthemeraux. Tethys, 5, 209-231. Macquart-Moulin, C. (19774. Les r6actions photocinetiques des peracarides du plancton nocturne. Fraction plantonique : Gmtrosaccua lobatua et Gmtrosaccua armatus. Tethya, 7, 1976, 339-348. Macquart-Moulin, C. (197713).Les modifications des rBactions photocinetiques des peracarides de l’hyponeuston nocturne en fonction de la composition spectrale de la lumi8re. Tethys, 7, 1975, 349-356. Macquart-Moulin, C. (1977~).Le controle de l’emergence et des nages nocturnes chez les peracarides des plages de MBditerranBe. Eurydice a . n k Hansen (Isopoda). Gmtroaaccua mediterraneua Bacescu, Gastrosaccua spinifer (Goes) (Mysidacea). Journal of Experimental Marine Biology and Ecology, 27, 61-81. Macquart-Moulin, C. and Leveau, M. (1968). RBcolte de macroplancton iL partir de la b o d e laboratoire-juin 1966. Rapport et procea-verbaux des rduniona. Commission internationale pour I’exploration acientifiue de la mer Mdditerranke, 19,495-497.

REFERENCES

293

Macquart-Moulin,C. and Patriti, G. (1966).Remarque sur la biologie d'Hemimyais speluncola Ledoyer. Mysidace sciaphile des grottes sous-marines obscures de la region de Marseille. Recueille des Travaux de la Station Marine d'Endoume, Facultd-deaSciences de Marseille, 40, 253-258. Makings, P. (1977).A guide to British coastal Mysidacea. Field Studies, 4,575-595. Makings, P. (1978). Two rare species of Schistomyeis (Mysidacea) from South Wales. Cmtaceana, 35, 223-224. Mankowski, W. (1962). Macroplankton as indicator of saline idlux of North Sea waters into the Baltic. Annabs Biologiques, 17, (1960), 90. Manton, S. M. (1928a). On some points on the anatomy and habits of the lophogastrid Crustacea. Transactions of the Royal Society of Edinburgh, 56, 103-119. Manton, S. M. (1928b). On the embryology of a mysid crustacean, Hemimysis lantornaa. Philosophical Transactions of the Royal Society, B , 216, 363-463. Markle, D. F. and Grant, G. C. (1970). The summer food habits of young-of-theyear striped bass in three Virginia rivers. Chesapeake Science, 11, 50-54. Marsh, G. A. (1973). The Zostera epifaunal community in the York River, Virginia. Chesapeake Science, 14, 87-97. Marshall, J. S., Waller, B. J. and Yaguchi, E. M. (1975). Plutonium in the Laurentian Great Lakes : food-chain relationships. Verhandlungen der Intenzationalen Vereinigung f iir theoretiache und angewandte Limnologie, 19, 323-329. Martinov, A. (1924). On some interesting Malacostraca from fresh-waters of European Russia. Ruaskii gidrobiologicheskii Zhurnal, 3, 210-216. Massonet, R. (1960). Carotenoides e t vitamine A chez les invertebres marins. Station de Aquiculture et & P b h e de Cmtiglione, 10, 137-173. Matsudaira, C., Kariya, T. and Tsuda, T. (1952). The study of the biology of a mysid Gmtrosaccua vulgaris Nakasawa. Tohoku Journal of Agricultural Research, 3, 155-174. Matthews, J. B. L. and Bakke, J. L. W. (1977). Ecological studies on the deepwater pelagic community of Korsfjorden, western Norway. The search for a trophic pattern. Helgolander wiwsemchaftliche Meeresuntermhungen, 30, 47-61. Mauchline, J. (1965). Breeding and fecundity of Praunwr inermie (Crustacea, Mysidacea).Journal of the Marine Biological Association of the United Kingdom, 45, 663-671. Mauchline, J. (1966). Thalmsomycesfagei, an ellobiopsid parasite of the euphausiid crustacean, Thysanoessa rmchi. Journal of the Marine Biological Association of the United Kingdom, 46, 531-539. Mauchline, J. (1967). The biology of Schietmyeis spirituS (Crustacea, Mysidacea). Journal of the Marine Biological Association of the United Kingdom, 47, 383-396. Mauchline, J. (1968). The biology of Erythrops serrata and E. ekgans (Crustacea, Mysidacea).Journal of the Marine Biological Association of the United Kingdom, 48, 455-464. Mauchline, J. (1969a). The biology of LeptomyG gracilb and L. lingvura (Crustacea, Mysidacea). Journal of the Marine Biological Association of the United Kingdom, 49, 379-389. Mauchline, J. (196913).Choniostomatid parasites on species of Erythrops (Crustacea, Mysidacea).Journal of the Marine Biological Association of the United Kingdom, 49, 391-392.

294

THE BIOLOGY OF MYSIDS

Mauchline, J. (19704. The biology of Schistomysis ornata (Crustacea, Mysidacea). Journal of the Marine Biological Association, of the United Kingdom, 50, 169-175. Mauchline, J. (1970b). The biology of Mysirlopeis gibbosa, M . didelphys and A!. angusta (Crustacea, Mysidacea). Journal of the Marine Biological Association of t h United Kingdom, 50, 381-396. Mauchline, J. (19714. The biology of Paramysis arelaosa (Crustacea, Mysidacea). Journal of the Marine Biological Association of the United Kingdom, 51, 339-345. Mauchline, J. (1971b). The biology of Neomysis integer (Crustacek, Mysidacea). Journal of the Marine Biological Association of the United Kingdom, 51, 347-354. Mauchline, J. (1971~).The biology of P r a u n w jlexuosw, and P. neglectwr (Crustacea, Mysidacea). Journal of the Marine BioEogicaE Association of the United Kingdom, 51, 641-652. Mauchline, J. (1971d). The biology ofSchistomysis kervilki (Crustacea, Mysidacea). Journal of the Marine Biological Association of the United Kingdom, 51, 653-658. Mauchline, J . (19718).Rare species of Mysidacea (Crustacea) from the west coast of Scotland. Journal of the Marine Biological Association of the United Kingdom, 51, 799-808. Mauchline, J. (1971f). Seasonal occurrence of mysids (Crustacea) and evidence of social behaviour. Journal of the Marine Biological Association of the United Kingdom, 51, 809-825. Mauchline, J. (1971g). Crustacea: Mysidacea. I n “Fauna of the Clyde Sea Area”, pp. 1-26. Scottish Marine Biological Association. Mauchline, J. ( 1972).The biology of bathypelagic organisms, especially Crustacea. Deep-sea Research, 19, 753-780. Mauchline, J. (1973a). Intermoult growth of species of Mysidacea. Journal of the Marine Biological Association of the United Kingdom, 53, 569-572. Mauchline, J. (1973b). The broods of British Mysidacea (Crustacea). Journal of the Marine Biological Association of the United Kingdom, 53, 801-817. Assessing similarity between samples of plankton. J o u r d Mauchline, J. (1973~). of the Marine Biological Association of India, 14 (1972), 26-41. Mauchline, J. (1976). The Hiatt growth diagram for Crustacea. Marine Biology, 35, 79-84. Mauchline, J. (1977s). The integumental sensilla and glands of pelagic Crustacea. Journal of the Marine Biological Association of the United Kingdom, 57, 973-994. Mauchline, J. (1977b). Growth of shrimps, crabs and lobsters-an assessment, Journal du Conaeil, 37, 162-169. Mauchline, J. (1977~). Growth and moulting of Crustacea, especially euphausiids. I n “Oceanic Sound Scattering Prediction” (Eds N. R. Andersen and B. J. Zahuranec), Marine Science, Vol. 5, pp. 401-422. Plenum Press, New York and London. Mauchline, J. and Fisher, L. R. (1969). The biology of eupheusiids. Advance8 in Marine Biology, 7, 1-454. Mauchline, J. and Murano, M. (1977). World list of the Mysidacea, Crustacea. Journal of Tokyo University of Fisheries, 64, 39-88. Mauchline, J. and Nemoto, T. (1977). Integumental sensilla of diagnostic value in euphausiids. Journal of the Oceanographic Society of Japan, 33, 283-289.

REFEREROES

296

Mauchline, J., Aizawa, Y., Ishimaru, T., Nishida, S. and Marumo, R. (1977). Integuments1 sensilla of pelagic decapod crustaceans. Marine Biology, 43, 149-155. Maurer, D., Kinner, P., Leathem, W. and Watling, L. (1976)..Benthic faunal assemblages off the Delmarva Peninsula. Estuarine and Coastal Marine Science, 4, 163-177. Maynard, S. D., Riggs, F. V. and Walters, J. F. (1975).Mesopelagic micronekton in Hawaiian waters: faunal composition, standing stock, and die1 vertical migration. Fishery Bulletin, National Oceanic Atmsphe& Adminiatration of tht. United States, 73, 726-735. Mayrat, A. (1956).Mise en Bvidence de tendons chez les crustacBs dans le muscle attracteur du sinciput de Praunua jlexuoeua 0. F . Miiller. Bulletin de la Societd Zoologique de France, 80, 81-85. Mayrat, A. (1956a).Oeil, centres optiques et glandes endocrines de Praunua jlexuoswr (0.F . Miiller) (CrustacBs MysidacBs). Archives de Zoologie Expdrintentale et Qdndrale, 93, 319-366. Mayrat, A. (195613).Le systhme artBrie1de PraunuaJEemsuset le pretendu coeur frontal des malacostrac6s. Bulletin de la Station Ockanographiquede SalammbG, No. 53, 44-49. Mayrat, A. (1962).Premiers resultats d’une Btude au microscope Blectronique des yeux des crustac6s. Compte Rendu Hebdornadaire des Shnces de 1 ’ A d m i e des Sciences, 255, 766-768. Meek, A. (1901).Biological investigations. Report on the Scientific Investigations, Northumberland Sea Fisheries Committee, 26-30. Meek, A. (1903).Biological investigations. Report on the Seientifi Investigations, Northumberland Sea Fiaheries Committee, 25-26. Meek, A. (1908). Trawling experiments. Report on the Scientific Investigations, Northumberland Sea Fisheries Committee, 1907, 7-19. Meek, A. (1909).Surface and bottom life obtained a t the trawling experiments. Report on the Scientific Investigations, Northumberland Sea Fk3ht.rks Commitlee, 1908-1909, 27-29. Merrett, M. R. and Roe, H. S. J. (1974).Patterns and selectivity in the feeding of certain mesopelagic fishes. Marine Biology, 28, 116-126. Meusy, J. J. (1963). Description de la glande androgene chez d e w crustacBs PBracarides : Paramyeis nouveli Labat (MysidacB)et E o c u m dollfusi Calman (CumacB).Compte Rendu Hebdomadaire dea S k a w de 1 ’ A d m i e dea Sciences, 256, 5425-6428. Mickel, T.J. and Childress, J. J. (1978).The effect of pH on oxygen consumption and activity in the bathypelagic mysid Uruzthophuuak ingens. Biological Bulletin, Marine Biological Laboratory, Wood’s Hole, M a e . , 154, 138147. Mielck, W.(1911).Quantitative untersuchungen an dem Plankton der deutschen Nordsee-Terminfahrten im Februar und Mai 1906. Wiasenschaftliche Meeresuntersuchungen der Kommiesion Wksenschaftlkhen Untersuchung der deuhchen Meere, 12, 315-347. Mikhman, A. S. (1978).Nutrition and feeding requirements of the percarina Percarina demidofi Nordmann larvae in the eestern part of the Gulf of Taganrog. Voprosy Ikhtiologii, 18, 366-370. Mikhman, A. S. and Romanovich, L. V. (1977).On the feeding of the Azov anchovy Engraulis encrasicholua maeoticua Pusanov. Vorpo@ Ikhtiologii, 17, 270-274.

296

TEE BIOLOGY OF MYSIDS

Miller, B. S. (1967). Stomach contents of adult starry flounder and sand sole in East Sound, Orcas Island, Washington. Journal of the Fisheries Research Board of Canada, 24, 2515-2526. Miller, B. S. (1970). Food of the flathead sole (Hippoglossoides elmsodon) in East Sound, Orcas Island, Washington. Journal of the Fisheriea Research Board of Canada, 27, 1661-1665. Miller, J. M. (1973). A quantitative push-net system for transect studies of larval fish and macrozooplankton. Limnology and Oceanography, 18, 152-154. Microshnichenko,M. P. and Vovk, F. I. (1973). A model of the Mysidae production process as exemplified with Paramy& intermedia (Czern.) from Tsimlyanskoye Reservoir. CfidrobiologicheakiiZ h u m l , 9, 36-44. Moguilevsky, A. and Gooday, A. J. (1977). Some observations on the vertical distribution and stomach contents of Cfigantocypris mwlleri Skogsberg 1920 (Ostracoda,Myodocopina).Sixth International Ostracod Symposium, Saaljelden, 263-270.

Mohr, J. L. (1953). Some observations on Arctic crustaceans and their associates: notes on the fauna of Nuwak Pond, Point Barrow. I n “Current Biological Research in the Alaskan Arctic”, pp. 14-18. Stanford University Publications, University Series, Biological Sciences, Vol. XI. Molander, A. R. (1962). Studies of the fauna in the fjords of Bohuslan with reference to the distribution of different associations. Arkiv for Zoologi, Ser. 2, 15, 1-64.

Molenock, J. (1969). Mysidopsis bahia, a new species of mysid (Crustacea: Mysidacea)from Galveston Bay, Texas. Tulane Studies in Zoology and Botany, 15, 113-116.

MBller, H. (1979). Significance of coelenterates in relation to other plankton organisms. Meereaforschung, Reports on Marine Reaearch, 27, 1-18. Molloy, F. M. E. (1958). The comparative anatomy and histophysiology of the alimentary canal in certain Mysidacea. Thesis presented for the degree of Ph.D., University of London. Moore, E. and Sander, F. (1977). A study of the offshore zooplankton of the tropical western Atlantic near Barbados. Ophelia, 16, 77-96. Moore, H. B. (1937). Marine fauna of the Isle of Man. Proceeding8 and Transactions of the Liverpool Biological Society, 50, 1-293. Moore, J. W. (1976). The proximate and fatty acid composition of some estuarine crustaceans. Estuarine and Cowtat Marine Science, 4, 215-224. Moore, J. W. and Moore, I. A. (1976). The basis of food selection in flounders, Platichthys fieam (L.) in the Severn Estuary. Journal of Fish Biology, 9, 139-156.

Moore, J. W., Moore, I. A. and Claridge, P. N. (1979). Seasonal changes in density, composition and reproductive biology of crustacean populations in the Severn Estuary. Crwrtuceana, 86, 113-122. Moran, S. (1972). Ecology and distribution of the sand-dwellingmysid Gmtrouaccua sanaua (Van Beneden, 1861) along the Mediterranean sandy shore of Israel. Crwrtaceana, SUPP.3, 357-361. Moran, S. and Fishelson, L. (1971). Predation of sand-dwelling mysid crustacean Cfmtrosaccw, sanctwr by plover birds (Charadriidae). Marine Biology, 9, 63-64.

Mordukhai-Boltovskoi, P. D. (1979). Composition and distribution of Caspian fauna in the light of modern data. Interntionale Revue der gaamten Hydrobwlogie, 64, 1-38.

REFERENCES

297

Morgan, M. D. and Beeton, A. M. (1978). Life history and abundance of Mysis relic& in Lake Michigan. Journal of the Fisheries Research Board of Canada, 85, 1165-1170.

Morris, R. J. (1971). Seasonal and environmental effects on the lipid composition of Neomysis integer. Journal of the Marine Biological Association of the Unite& Kingdom, 51, 21-31. Morris, R. J. (1972). ‘ Q e occurrence of wax esters in crustaceans from the northeast Atlantic O c e p . Marine Biology, 16, 102-107. Morris, R. J. (1973). R9tionships between the sex and degree of maturity of marine crustaceans and their lipid compositions. J o u m l of the Marine Biological Association of the United Kingdom, 53, 27-37. Morris, R. J. and Sargent, J. R. (1973). Studies on the lipid metabolism of some oceanic crustaceans. Marine Biology, 22, 77-83. Morris, R. J., Ferguson, C. F. and Raymont, J. E. G. (1973). Preliminary studies on the lipid metabolism of Neomysis integer, involving labelled feeding experiments. Journal of the Marine Biological Association of the United Kingdom, 58, 657-664. Morris, R. J., Armitage, M. E., Raymont, J. E. G., Ferguson, C. F. and Raymont, J. K. B. (1977). Effects of a starch diet on the lipid chemistry of Neomysis integer (Leach). Journal of the Marine Biological Association of the United Kingdom, 57, 181-189. Morswell, J. W-. and Norden, C. R. (1968). Food habits of the alewife, Alosa pseudoharengw, (Wilson), in Lake Michigan. Proceedings of the Eleventh Conference on Great Lakes Research, 98-102. Mukai, H . (1971). T h e phytal animals on the thalli of Sargmsum serratifolium in the Sargassum region, with reference to their seasonal fluctuations. Marine Biology, 8, 170-182. Muncy, R. J. (1962).Life history of the yellow perch, Perca$avescem, in estuarine waters of Severn River, a tributary of Chesapeake Bay, Maryland. Chesapeake Science, 3, 143-159. Mundie, J. H. (1959). The diurnal activity of the larger invertebrates a t the surface of Lac la Rouge, Saskatchewan. Canadian Journal of Zoology, 37, 945-956. Murano, M. (1963a). Neomysis intermedia Czerniavsky. I Role in lake production. Japan Aquiculture Research Group, 11, 149-158. Murano, M. (1963b). Neomyk8 intermedia Czerniavsky. 11. Food and feeding habits. Japan Aquiculture Research Group, 11, 159-165. Murano, M. (19644. Fisheries biology of a marine relict mysid Neomysi.9 intermedia Czerniavsky. 111. Lifecycle, with special reference to the reproduction of the mysid. Japan Aquiculture Research Group, 12, 19-39. Murano, M. (1964b).Fisheries biology of a marine relict mysid Neomysis intermedia Czerniavsky. N. Lifecycle with special reference to growth. Japan Aquiculture Research Group, 12, 109-117. Murano, M. (1966a). Two new species of Pseudomma (Mysidacea) from Sagami Bay, Central Japan. Journal of the OceanographicalSociety of Japan, 22,41-49. Murano, M. (1966b).Descriptions of two new species of Heteroerythrops (Mysidacea) from Japan. Journal of the Oceanographical Society of Japan, 22, 111-118. Murano, M. (1966~).Culture of I s a z a - h i Neomysis intermedia. Information Bulletin of Planktology in Japan, 13, 65-68. Murano, M. (1966d). Neomysis intermedia Czerniavsky. V. Adaptability to environment. Japan Aquiculture Research Group, 13, 233-245.

298

THE BIOLOGY OF MYSIDS

Murano, M. (19668). Neomysis intermedia Czerniavsky. VI. Possibility of transference from brackish to fresh water. Japan Aquiculture Research aroup, 14, 79-84. Murano, M. (1969). Three new species of Mysidacea from Japan. Cmtaceana, 17, 207-21 9. Murano, M. (1970a).Three species belonging to the genus Lophogaster (Mysidacea) from Japan. Proceedings of the Japanese Society of Systematic Zoology, No. 6, 1-5. Murano, M. (1970b). A small collection of benthic Mysidacea from coastal waters in Suruga Bay, Japan. Cmtaceana, 18, 251-268. Murano, M. ( 1 9 7 0 ~ )Systematic . and ecological studies on Mysidacea collected by the bottom net. Journal of the Oceanographical Society of Japan, 26, 137-150. Murano, M. (1974a). Scolamblyops japonicwr gen. nov. sp. nov. (Mysidacea) from Suruga Bay, Japan. Crwrtaceana, 26, 225-228. Murano, M. (1974b). Mysidacea from the central and western Pacific. I. Genus Pseudomma (Tribe Erythropini). Publications of the Seto Marine Biological Laboratory, 21, 287-334. Murano, M. (1975). Mysidacea from the central and western Pacific. 11. Hyperamblyops, Teraterythrops and Synerythrops (Tribe Erythropini.) Publications of the Seto Marine Biological Laboratory, 22, 81-103. Murano, M. (1976). Mysidacea from the central and western Pacific. 111. Genera Eoerythrops, Holmiella, Fterornysis, Longithorax and Katerythrops (Tribe Erythropini). Publications of the Set0 Marine Biological Laboratory, 23, 19-50. Murano, M. (1977a).Mysidacea from the central and western Pacific. IV. Genera Euchaetomera, Euchaetomeropsis, Arachnomysis, Caesarmysis, Echinomysides, Meterythrops and Nipponerythrops (Tribe Erythropini). Publicatiom of the Set0 Marine Biological Laboratory, 24, 141-192. Murano, M. (1977b). Five new species belonging to the genus Proneomysis (Crustacea, Mysidacea) from Japan. Bulletin of the National Science Mwreurn of Japan, Series, A , Zoology, 3, 225-240. Murano, M. (1978).A new species of Idiomysb (Crustacea, Mysidacea) from Japan. Bulletin of the National Science M w e u m , Series A , Zoology, 4, 263-266. Murano, M., Marumo, R., Nemoto, T. and Aizawa, Y. (1976).Vertical distribution of biomass of plankton and micronekton in the Kuroshio water off central Japan. Bulletin of the Plankton Society of Japan, 23, 1-12. Muus, B. J. (1967). The fauna of Danish estuaries and lagoons. Meddelelser fra Danmarks Fiskeri-og Havundersogelser, N.S., 5 (l),1-316. Nagabhushanam, A. K. (1965). On the biology of the commoner gadoids in Manx waters. Journal of the Marine Biological Association of the United Kingdom, 45, 615-657. Nair, K. B. (1939). The reproduction, oogenesis and development of Mesopodopsb orientalis Tatt. Proceedings of the Indian Academy of Sciences, 9, 175-223. Nakamura, E. (1967). Abundance and distribution of zooplankton in Hawaiian waters, 1955-56. Special Scientijic Report, United States Department of the Interior, Fish and Wildlqe Service, No. 544, 1-37. Nakazawa, K. (1910). Notes on Japanese Schizopoda. Annotationes ZooZogiCae japonenses, 7, 247-261. Nath, C. N. (1972). On the storage of calcium in Spebeomysis, a subterranean mysid. Cwtaceana, Supp. 3, 351-353. Nath, C. N. (1973). Breeding and fecundity in a subterranean mysid, Lepidomysis longipes (Pillai and Mariamma). International Journal of Speleology, 5,319-323.

REFERENCES

299

Nath, C. N. (1974).Studies on the abdominal musculature of the subterranean mysid Lepidomyais longipea (Pillai and Mariamma). International Journal of Speleology, 6, 173-180. Nath, C. N. and Pillai, N. K. (1971).Studies on the genus Lepidomysis Clarke. Crustacea : Mysidacea. Hydrobiologia, 37, 285-300. Nath, C. N. and Pillai, N. K. (1972).On the alimentary canal of S p e h m y G longipes (Crustacea: Mysidacea).Journal of the ZooZogiCc2.1Society of India, 23, 1971, 95-108. Nath, C. N. and Pillai, N. K. (1976).The alimentary system of the littoral mysid Gaatrosaccus sinaulans (Van Beneden). Journal of the Marine Biological Association of India, 15, 1973, 577-586. Nath, C. N., Thampy, D. M. and Pillai, N. K.(1972).A note on the androgenic gland of a subterranean mysidacean, Spelaeohysia longipea (Pillai and Mariamma). Crustaceam, Supp. 3, 354-356. Needham, A. E. (1937).Some points in the development of NeomysL vulgaris. Quarterly Journal of Microscopical Science, 79, 559-588. Neil, D. M. (1975a).The control of eyestalk movements in the mysid shrimp Praunus j l e w a u s . Journal of E e r i m e n t a l Biology, 62, 487-504. Neil, D. M. (1975b).The optokinetic responses of the mysid shrimp Praunus jlexuosua. Journal of Experimental Biology, 62,505-518. Neil, D.M. (1975~). The mechanism of statocyst operation in the mysid shrimp Praunus jlexuoaus. Journal of Experimental Biology, 62, 685-700. Nekrasova, M. Ye. and Rakitina, N. I. (1968). On ratio of weight and linear dimensions in mysids of T a g m o g Bay. Gidrobiologicheekii Zhurnal, 4,52-54. Nemoto, T. (1959).Food of baleen whales with reference to whale movements. ScientiJic Reports of the Whales Reaearch Imtitute, Tokyo, No. 14, 149-290. Nemoto, T. (1972). Chlorophyll pigments in the stomach and gut of some macrozooplankton species. I n “Biological Oceanography of the Northern Pacific Ocean” (EdsA. Y. Takenouti et aZ.), pp. 411-418. Idemitsu Shoten, Tokyo. Nicol, J. A. C. (1959).Studies on luminescence. Attraction of animals t o a weak light. JownaZ of the Marine BioZogicaZ Association of the United Kingdbm, 88, 477-479. Nikolaev, I. I., Andronikova, I. N. and Shushkina, E. A. (1973).On the vertical distribution of zooplankton (excluding protozoans) in the deep-water region of Lake Ladoga. Translation Series, Fisheriea Reaearch Board of Canada, No. 2848, 1-6. Nimmo, D. R., Rigby, R. A., Bahner, L. H. and Sheppard, J. M. (1978).The acute and chronic effects of cadmium on the estuarine mysid My&opG bahia. Bulletin of Environmental Contarnination and Toxicology, 19, 80-85. Norman, A.M. (1865).Reports on deep sea dredging on the coasts of Northumberland and Durham. Crustacea. Natural History Transactions of Northumberland and Durham, 1, 12-29. Norman, A. M. (1880a). Notes on the French explorations of “Le Travailleur” in the Bay of Biscay. Annals and Magazine of Natural History, Ser. IT, 430-436. Norman, A. M. (1880b). Report on the Crustacea. Proceedings of the Royal Society of Edinburgh, 1881-1882, 683-689. Norman, A. M. (1886).On a Crangon, some Schizopoda and Cumacea new to, or rare in British seas. Annals and Magazine of Natural History, Ser. V, 19, 89-103.

300

THE BIOLOGY OF MYSIDS

Northcote, T. G. (1972).Kootenay Lake: man’s effects on the salmonid community. Journal of the Fisheries Research Board of Canada, 29, 861-865. Northcote, T. G. (1973). Some impacts of man on Kootenay Lake and its salmonids. Great Lakes Fishery Commission, Technical Report, No. 25, 1-46. Nouvel, H. (1937). Observation de l’accouplement chez une espbce de Mysis, Praunus jlexuosus. Co&pte Rendu Hebdomadaire des Shnces de l’dcadernie des Sciences, 205, 1184-1 186. Nouvel, H. (1940). Observations sur la sexualit6 d’un mysidac6, Heteromysis armoricana n.sp. Bulletin de Mw’& Ocdanographiquede Monaco, No. 789, 1-11. Nouvel, H. (1941). Sur les ellobiopsides des mysidaces provenant des Campagnes du Prince de Monaco. Bulletin de Muade Ockanographique de Monaco, No. 809, 1-8. Nouvel, H. (1942). Diagnoses preliminaires de mysidaces nouveaux provenant des Campagnes du Prince Albert I de Monaco. Bulletin de Muske Ochnographique de Monaco, No. 831, 1-12. Nouvel, H. (1944a). Diagnoses de mysidac6s nouveaux de la Mer Rouge et du Golfe d’Aden. Bulletin de la Socidtd d’Histoire Naturelle de Toulouse,. 79, 255-269. Nouvel, H. (1944b). L’inversion de l’asymetrie chez les crustaces avec description d’un nouveau cas observe chez des Processa. Bulletin de l’lnstitut Ochnographique, No. 865, 1-6. Nouvel, H. (1945). Les relations entre le periodicit6 lunaire, les marees et la mue des crustads. Bulletin de I’Institut Ocdanographique, No. 878, 1-4. Nouvel, H. (1950). Mysidacea. Conseil International pour I’Explorution de la Mer, Zooplancton Fiche, Nos 18-27. Nouvel, H. (1951a). Les mysidaces des cdtes du Maroc. Bulletin de la SocidtB des Sciences Naturelles du Maroc, 31, 37-40. Nouvel, H. (1951b). Gmtrosaccus normani G. 0. Sars 1877 et Gmtrosaccus lobatus, n.sp. (Crust. Mysid.) avec precision de l’hbte de Prodajus lobiuncoi Bonnier (Crust, hop. Epicar.). Bulletin de Mmde Ocdanographiquede Monaco, No. 993, 1-12. Nouvel, H. (1952). La Mysis frontalis de Milne Edwards. Bulletin du Museum National d’Histoire Naturelb, Ser. 2, 24, 519-521. Nouvel, H. (1953). Un Ellobiopsidae nouveau (AmallocystG boschmai nap.) parasite d’un mysidac6 en MBditerranee (Note pr6liminaire). Vie et Milieu, 4, 57-58. Nouvel, H . (1954). Description d‘un mysidace nouveau de Ceylon Mesopodopsis zeylanica n.sp. Zoologische Mededeelingen, 33, 33-39. Nouvel, H. (1957a). Mysidaces provenant de deux Bchantillons de “Djembret” de Java. Zoologische Mededeelingen, 35, 315-331. Nouvel, H. (195713). L’exuviation chez les Mysidae (CrustacBs, Mysidaces). Bulletin de la Socidtd zoologique de France, 82, 395-400. Nouvel, H. (1960). Presence en MBditerranee d’un Mysidae de la sous-famille des Rhopalophthalminae : Rhopalophthalmus mediterraneus n.sp. Bulletin de la Sociktk d’Histoire Naturelle de Toulouae, 95, 225-231. Nouvel, H. (1964a). MysidacBs recoltes par S. Frontier iL Nose-Be. I. Description de deux Leptomysini. Bulletin de la Socidtd d’Histoire Naturelle de Toulouse, 99, 107-123. Nouvel, H. (1964b). Heterowaysis gerlachei (Bonnier and Perez) et les esp6ces actuellement confondues SOUS le nom d‘Heteromyeis harpax (Crustacea Mysidacea). Zoologische Mededeelingen, 39, 37-44.

REFERENUES

301

Nouvel, H. (1964~). RBgBnBrations hethomorphiques de 1’Bcaille antennaire chez les mysidacBs. Vie et Milieu, Supp. 17, 337-344. Nouvel, H. (1965). MysidacBs r6coltes par S. Frontier B Nosy-BB. 11. Description de deux Mysini appartenant aux genres Diamysis et Acanthomysis. Bulletin de la Socidtd d’Histoire Naturelle de Toulouse, 100, 451-464. Nouvel, H. (1966a). Les genres Tenagomysis Thompson 1900, Doxomysis Hansen, 1912, AfroFysis Zimmer, 1916 et Iimysis n.gen. (Crust. Mysidacea Leptomysini). Bulletin de la Socidtd d’Histoire Naturelle de Toulouse, 102, 319-324. Nouvel, H. (196613). Mysidaces r6coltBs par S. Frontier B Nosy-BB. 111. HyperiiTnyS’i8 daga-scariensis n.gen., n.sp., Leptomysini presentant une singulidre modification des appendices thoraciques de la dernidre paire. Bulletin de la Socidtd &Histotoire de Toulouae, 102, 493-505. Nouvel, H. (1967).Mysidaces rBcoltBs par S. Frontier B Nosy-BB. IV. Mesacanthomy& pygmaea n.gen., n.sp., et Anisomysis hanseni, n.sp. Bulletin de la Socidtd d’Hktoire Naturelle de Toulouse, 102, 493-505. Nouvel, H. (1969). MysidacBs rBcolt6s par S. Frontier B Nosy-BB. V. Anchialina n.sp. Bulletin de la Socidtd d’Histoire Naturelle de Toulouse, &aqmcarie& 105, 340-356. Nouvel, H. (1971). MysidacBs rdcoltes par S. Frontier it Nosy-BB. VI. Anchhlina typka (Kroyer) avec distinction de deux sous-espdces et Anchialina latqrons n.sp. Bulletin de la Socidtd d’Histoire Naturelle de Toulouse, 107,325-338. Nouvel, H. (1973a). Pseudoxomysis caudaensis n.gen., n.sp., crustace mysidacB (Leptomysini) de la Mer de Chine meridionale. Bulletin de la Socidtd d’Histoire Naturelle de Toulouse, 109, 131-141. Nouvel, H. (1973b). Une mysidacB de la Nouvelle-Caledonie: A n i s o m y G thurneysseni n.sp. Bulletin du Museurn Nat,ional d’Histoire Naturelle, Ser. 3, 185, (Z0010gie 124), 1453-1459. Nouvel, H. (1973~).Observations sur les mysidacBs et quelques cumaces littoraux de lct c6te francaise du Golfe de Gascogne au sud de l’embouchure de la Gironde. Bulletin du Centre d’Etudeaet de Recherches Scienti$quee, Biarritz, 9, 1972, 127-140. Nouvel, H. (1973d). Sur quelques mysidacBs de 1’Afrique du Sud avec description d’une espdce nouvelle du genre Erythrops. Bulletin de la Socidtd d’Histoire Naturdle de Toulouse, 109, 241-247. Nouvel, H. (1974).Mysidaces rBcoltBs par S. Frontier B Nosy-BB. VII. Description de deux Erythropini nouveaux appartenant aux genres Erythrops et Hyperelythrops. Bulletin de la Socidtd d’Hktoire Naturelle de Toulouse, 110, 7-20. Nouvel, H. (1978).Mysidaces rBcoltBs par S. Frontier a Nosy-BB. VIII. Lophogaster afinis Colosi et Mesopodopsis africana daga-scariensis ssp. nov. Bulletin de la SociWd d’Histoire Naturelle de Toulouse, 113, 396-398. Nouvel, H. and Hoenigman, J. (1955). Amallocystk boschmai Nouvel, 1954, ellobiopside parasite du mysidacB Leptomysis gracilis G. 0. Sars. Vie et Milieu, SUPP. 6, 1-18. Nouvel, H . and Lagarddre, J. P. (1976). Les mysidacBs du talus continental du Golfe de Gascogne. I. Tribu des Erythropini (genre Eqthrops except6). Bulletin d u Museum National #Histoire Naturelle, Ser. 3, 414 (Zoologie, 291), 1243-1324. Nouvel, H. and Nouvel, L. (1939). Observations sur la biologie d‘une Mysis: Praunus Jexuosus (Miiller, 1788). Bulletin du Mus& Ockanographique de Monaco, No. 761, 1-10.

302

THE BIOLOQY OF MYSIDS

Nouvel, H. and Panouse, J. B. (1965). Presence de Paramysis nouveli (Mysidacea) et de Crangon crangon (Decapoda Natantia) (Crustacea) sur la cBte Atlantique du Maroc. Bulletin de la Socie'ti des Sciences Naturellea.et Physiques du Maroc, 45, 29-31. Nuorteva, P. and Hiisiinen, E. (1975). Bioaccumulation of mercury in Myozowphalus quadricornis (L.) (Teleostei, Cottidae) in an unpolluted area of the Baltic. Annales Zoologici Fennici, 12, 247-254. Nusbaum, J. and Schreiber, W. (1898). Beitrage zur Kenntnis der sogenannten Ruckenorgane der crustaceen Embryonen. Biologischea Zentralblatt, 18, 136-746. Nusbaum, K.(1887). L'embryologie de Mysis chameko (Thompson). Archivee de Zoologie Experimentale et Odne'rale, Series 2, 5, 123-202. Nyman, L. and Westin, L. (1969). A contribution to the methods of classification of some Mysidae and Gammarw species in the Baltic. Report of the Institute of Freshwater Research, Drottni~ghoZm,49, 157-163. Ogawa, Y . (1964). A new record of Paracanthomyais hispida I i from the Japan Sea. Bulletin of the Japan Sea Regional Fisheries Research Laboratory, 13, 131-132. Ogle, J. and Price, W. (1976). Growth of the shrimp, Penaew aztecw, fed a diet of live mysids (Crustacea: Mysidacea). Gulf Research Reports, 5, 46-41. Okada, S. and Taniguchi, A. (1971). Size relationships between salmon juveniles in shore waters and their prey animals. Bulletin of the Faculty of Fisheries, Hokkaido University, 22, 30-36. Omori, M. (1969a). Weight and chemical composition of some important oceanic zooplankton in the North Pacific Ocean. Marine Biology, 3, 4-10. Omori, M. (1969b). A bottom-net to collect zooplankton living close to the sea floor. Journal of the OceanographicalSociety of Japan, 25, 291-294. Omori, M. (1978). Zooplankton fisheries of the world: a review. Marine Biology, 48, 199-205. O'Riordan, C. E. (1969). A catalogue of the collection of Irish marine Crustacea in the National Museum of Ireland. Stationery Ofice,Dublin, 1-98. Ostenfeld, C. H. (1906). Catalogue des esphces de plantes et d'animaux observees dans la plankton recueilli pendant les expeditions periodiques depuis le mois d'eoiit 1902 jusqu'au mois de mai 1905. Publications de Circonstance, Con,seiE permanent international pour l'exploration de la m r , 33, 1-122. Ostenfeld, C. H. (1916). Catalogue des esphces de plantes et d'animaux observees dans le plankton recueilli pendant les expeditions depuis le mois juillet 1908 jusqu'au mois de decembre 191 1. Publications de Circonstance, Conseil permanent international pour l'exploration de la mer, 70, 1-81. Ostenfeld, C. H. and Wesenberg-Lund, C. (1909). Catalogue des esp6ces de plantes et d'animaux observees dam le plankton recueilli pendant les expeditions periodiques depuis le mois d'auot 1905 jusqu'au mois de Mai 1908. Publications de Circonstance, Conseil permanent international pour l'exploration de la rner, 48, 1-151. 0stvedt, 0.-J. (1955). Zooplankton investigations from Weather Ship M in the Norwegian Sea, 1948-49. Hvalradets Skrifter, No. 40, 1-93. Otto, J. P. and Wielinga, D. T. (1933). Hydrobiologische notizen vom brackwassergebeit der Provinz Friesland, speziell in der niihe von harlingen. Tijdschrift der Nederlandsche Dierkundige Vereeniging, 3, 49-14. Oug, E. (1977). Faunal distribution close to the sediment of a shallow marhe environment. SarSia, 63, 115-121.

REFERENOES

303

Overstreet, R. M. and Heard, R. W. (1978). Food of the Atlantic croaker, Micropogonias undulatus, from’ Mississippi Sound and The Gulf of Mexico. Gulf Research Reports, 6 , 145-152. Painter, R. E. (1966). Zooplankton of San Pablo and Suisun Bays. California Fish and Qm, 133, 18-39. Panampunnayil, S. U. (1977). Three new species of Mysidacea from Kerala coast. I n “Proceedings of the Symposium on Warm Water Zooplankton”, pp. 28-35. Special Publication of National Institute of Oceanography, Goa India. Parker, M. (1979). Neomysia integer (Leach) (Crustacea: Mysidacea): records and distribution in Ireland. Irish Naturalish’ Journal, 19, 339-342. Parker, M. and West, B. (1979). The natural history of Neomysia integer (Leach) in Lough Furnace, Co. Mayo, a brackish lough in the west of Ireland. Estuarine and CoaataE Marine Science, 8, 157-167. Pasternak, A. F. (1977). Changes in some energy transformation indices during the growth of the mysid Neomyais mirabilis (Czerniavsky). Oceanology, 17, 470-473.

Paulsen, 0. (1918). Plankton and other biological investigations in the sea around the Faeroes in 1913. Meddelelser f r a Kommkaionen for Havundersngelser, Serie Plankton, 1 (13), 1-27. Pawsey, E. L. and Davis, F. M. (1924). Report on exploratory voyages to Lousy Bank and adjacent areas. Fishery Investigations, Ministry of Agriculture, Food and Fisheries, Ser. 2, 7 (2), 1-22. Pearcy, W. G. and Ambler, J. W. (1974). Food habits of deep-sea macrourid fishes off the Oregon coast. Deep-sea Research, 21, 745-759. Pearcy, W. G., Krygier, E. E., Mesecar, R. and Ramsey, F. (1977). Vertical distribution and migration of oceanic micronekton off Oregon. Deep-sea Research, 24, 223-245. Pechen’-Finenko, G. A. and Pavlovskaya, T. V. (1975). Comparative evaluation of the role of detritus and algae in the nutrition of mysids Neomysis mirabilis (Czerniavsky).Qidrobiologischeskii Zhurnal, 11, 39-44. Pequegnat, L. H. (1965). The bathypelagic mysid Gmthophatmia (Crustacea) and its distribution in the eastern Pacific Ocean. Pacific Science, 19, 399-421. Peroival, E. (1929). A report on the fauna of the estuaries of the River Tamar and the River Lynher. Journal of the Marine Biological Association of the United Kingdom, 16, 81-108. Pesce, G. L. (1976a). A new locality for Spelaeomyai.9 botuzzii with redescription of the species (Crustacea, Mysidacea). Bollettino del Museo Civic0 d i Stoha Naturale d i Verona 11, 345-364. Pesce, G. L. (1975b). On a Stygiomysk (Crustacea, Mysidacea) from the southern Italy. Bollettino deZ Museo Civic0 d i Storia Naturale d i Verona, 11, 439-443. Petersen, C. G. J. (1914). Valuation of the Sea 11.The animal communities of the sea bottom and their importance for marine zoogeography. Report of the Danish Biological Station to the Board of Agriculture, 21, 1913, 1 4 4 ; Appendix 1-68.

Pezzack, D. S. and Corey, S. (1979). The life history and distribution of Neomysis americana (Smith): Crustacea, Mysidacea. Canadian Journml of Zoology, 57, 785-793.

Pillai, N. K. (1961). Additions to the Mysidacea of Kerala. Bulletin of the Research Institute of the University of Truvancore, 8, 15-35. Pillai, N. K. (1963a). On a new mysid from the inshore waters of the Kerala coast. Journal of the Marine Biological Association of India, 5 , 268-262.

304

THE BIOLOGY OF MYSIDS

Pillai, N. K. (196310). Description of a new species of Bathymyais with a revised definition of the genus. Journal of the Marine Biological Association of India, 5, 263-268.

Pillai, N. K. ( 1 9 6 3 ~ ) .Isopod parasites of Indian mysids. Annals and Magazine of Natural History, Ser. 13, 6 , 739-743. Pillai, N. K . (1963d). Redescription of Dioptromyeis perspicillata Zimmer. Bulletin of the Department of Marine Biology and Cccurwgraphy, University of Kerala. 1, 9-14. Pillai, N. K. (1964). Report on the Mysidacea in the collection of the Central Marine Fisheries Research Institute, Mandapam Camp, South India. Journal of the Marine Biological Association of India, 6 , 1-41. Pillai, N. K. (1965). A review of the work on the shallow water Mysidacea of the Indian waters. I n “Symposium on Crustacea”, 5, pp. 1681-1728. Marine Biological Association of India. Pillai, N. K. (1967). Description of the female of Mysidopsis indica W. M . Tattersall. Journal of the Marine Biological Association of India, 9, 121125.

Pillai, N. K. (1968). Redescription of Petalophthalmw o c u h t w (Crustacea: Mysidacea) based on an adult female collected near the type locality. Journal O f Zoology, 155, 283-291. Pillai, N. K. (1973). Mysidacea of the Indian Ocean. I n “Handbook to the International Zooplankton Collections”, 4, pp. 1-125. Indian Ocean Biological Centre, Kerala State, India. Pillai, N. K. and Mariamma, T. (1964). On a new lepidomysid from India. Cwtaceana, 7 , 113-124. Pillai, R. S. (1965). The circulatory system of Caridina ZaeG Heller. Cwtaceana, 8, 66-74,

Pitt, T. K. (1973). Food of American plaice (Hippoglossoides phtessoides) from the Grand Bank, Newfoundland. Journal of the Fisheries Research Board of Canada, 30, 1261-1273. Poirier, L. (1969). Distribution horizontale et verticale des mysidacds de la Baie des Chaleurs en 1968. Rapport, Station de Biologie Marine de Grade-Rivikre, 67-76.

Poirier, L., Granger, O., Weinstein, M. and Brunel, P. (1969). P h h e s hyperbenthiques et planctoniques quantitatives a l’entrde de la Baie des Chaleurs: protocole et methodes en 1968. Rapport, Station de Biologie Marine de QrandeRivikre, 47-57. Ponomarenko, I. Y . (1976). Food composition and quantitative indices of feeding of 0-group cod in the Barents Sea. Annales Biologiques, 31, (1974), 83-85. Ponomarenko, I. Y . (1978). Investigations on feeding and biological indices of 0-group cod in the Barents Sea. AnnaZes Biologiques, 33, (1976). 81-82. Powers, C. F. and Robertson, A. (1968). Biological exploration in Lake Michigan from a research submarine. Proceedings of the Eleventh Conference on Qreut Lakes Research, 117-123. Pratap, M. M., Wafar, M. V. M., Haridas, P., Narttyanan, B., Menon, P. G. and Sivadas, P. (1977). Comparative studies on the abundance of zooplankton in the surrounding sea and lagoons in the Lakshadweep. Indian Journal of Marine Sciences, 6 , 138-141. Prefontaine, G. and Brunel, P. (1962). Liste d’invertebrds marins recueilles dans l’estuaire du Saint-Laurent de 1929 21 1934. Le Naturaliste Canadien, 89, 237-263.

REFERENUES

305

Price, K. S. (1962). Biology of the sand shrimp, Crangon septemspinosa, in the shore zone of the Delaware Bay’region. Chesapeake Science, 3, 244-255. Price, W. W. (1975). A new locality record for Metamy&op8is swifti Bacescu, 1969 (Crustacea: Mysidacea) from the Gulf Coast of Mexico. Southwestern Naturalist, 20, 130. Price, W. W. (1978). Occurrence of Mysidopsis almyra Bowman, M . bahia Molenock and Bowlnanielh brmiliensis Bacescu (Crustacea, Mysidacea) from the eastern coast of Mexico. Gulf Research Reports, 6, 173-175. Price, W. W. and Vodopich, D. S. (1979). Occurrence of Mysidopsis almyra (Mysidacea, Mysidae) on the east coast of Florida, U.S.A.Crustuceana, 36, 194-196. Prygunkova, R. V. (1974). Certain peculiarities in the seasonal development of zooplankton in the Chupa Inlet of the White Sea. .ln “Explorations of the Fauna of the Seas. XI11 (XXI). Seasonal phenomena in the life of the White and Barents Seas, pp. 4-56, Zoological Institute, Academy of Sciences of the USSR. Pnibram, H. (1901). Experimentelle Studien uber Regeneration. Archiv f u r Entuvicklungsmechanik der Organismen, 11, 321-345. Quetin, L. B., Mickel, T. J. and Childress, J. J. (1978). A method for simultaneously measuring the oxygen consumption and activity of pelagic crustaceans. Comparative Biochemistry and Physiology, 59A, 263-266. Quintero, C. and Roa, E. Z. de (1973). Notas bioecologicas sobre Metamydopsis insularis Brattegard (Crustacea-Mysidacea) en una laguna litoral de Venezuela. Acta BioEogica Venezuelica, 8, 245-270. Quintero,C. and Roa, E. Z. de (1977). Misidaceos de algunas regiones de Manglares de Venezuela. Acta Biologica Venezuelica, 9, 259-270. Radtke, L. D. (1966). Distribution of smelt, juvenile sturgeon, and starry flounder in the Sacramento-SanJoaquin Delta with observations on food of sturgeon. California Fish and G a m , 136, 115-129. Randall, J. E., Schroeder, R. E. and Stark, W. A. (1964). Notes on the biology of echinoid D i a d e m antilhrum. Caribbean Journal of Science, 4, 421-433. Raw, F. (1955). The Malacostraca: their origin, relationships and phylogeny. Annals and Magazine of Natural History, Ser. 12, 8, 731-756. Rawson, D.S. (1942). A comparison of some large alpine lakes in western Canada. Ecology, 23, 143-161. Rawson, D. S. (1960). A limnological comparison of twelve large lakes in northern Saskatchewan. Lirnnology and Oceanography, 5,195-21 1. Rawson, D. S. (1961). The lake trout of Lac la Rouge, Sascatchewan. Journal of the Fisheries Research Board of Canada, 18, 423-462. Ray, R. (1914). Report on the examination of marine deposits in relation to the contents of fishes’ stomachs from the Irish Sea. Transactions of the fiverpool Biological Society, 28, 340-368. Raymont, J. E. G. (1961). The fauna of an inter-tidal mud flat. I n “Papers in Marine Biology and Oceanography”, pp. 170-203. Pergamon Press, Oxford. Raymont, J. E. G. and Conover, R. J. (1961). Further investigations on the carbohydrate content of marine zooplankton. fimnology and Oceanography, 6, 154-164. Raymont, J. E. G. and Krishnaswamy, S. (1960). Carbohydrates in some planktonic animals. Journal of the Marine Biological Association of the United Kingdom, 39, 239-240.

306

THE BIOLOGY OF MYSIDS

Raymont, J. E. G. and Krishnaswamy, S. (1968). A method for determining the oxygen uptake and carbon dioxide output in Neomyeia integer. Internationak Revue der Gesamten Hydrobiologie u. Hydrographie, 53, 563-572. Raymont, J. E. G. and Linford, E. (1966). A note on the biochemical composition of some Mediterranean zooplankton. Internationale Revue der GeaamtenHydrobiologie u. Hydrographie, 51,485-488. Raymont, J. E. G., Austin, J. and Linford, E. (1964). The biochemical composition of Neomysia integer. Journal du Conseil, 28, 354-363. Raymont, J. E. G., Austin, J. and Linford, E. (1966). Biochemical studies on marine zooplankton. 111. Seasonal variation in the biochemical composition of Neomysis integer. I n “Some Contemporary Studies in Marine Science” (Ed. H. Barnes), pp. 597-605. George Allen and Unwin Ltd., London. Raymont, J. E. G., Krishnaswamy, S. and Tundisi, J. (1967). Biochemical studies on marine zooplankton. IV. Investigations on succinic dehydrogenase activity in zooplankton with special reference t o Neomysis integer. Journal du Conseil, 31, 164-169. Raymont, J. E. G., Austin, J. and Linford, E. (1968). Biochemical studies on marine zooplankton. V. The composition of the major biochemical fractions in Neomysis integer. Journal of the Marine Biological Association of the United Kingdom, 48, 735-760. Raymont, J.E. G.,Srinivasagam,R.T.andRaymont,J.K. B. (1969).Biochemical studies on marine zooplankton. VII. Observations on certain deep sea zooplankton. Internationale Revue der GeaamtenHydrobiologie u. Hydrographie, 54, 357-365.

Raymont, J. E. G., Ferguson, C. F. and Raymont, J. K. B. (1973). Biochemical studies on marine zooplankton. The amino acid composition of some local species. Special Publication of the Marine Biological Association of India, 60, 91-99.

Redeke, H. C. (1941). Over het voedsel van eenige Ijsselmeer-Visschen. De Biologie van de Zuiderzee T i j h n s haar Drooglegging, 5, 27-56. Rees, C. B. (1939). The plankton in the upper reaches of the Bristol Channel. Journal of the Marine BiologicalAssociation of the United Kingdont,2a, 397-426. Reger, J. and Fain-Maurel, M. A. (1973). Studies on the origin, distribution and composition of the extracellular tubules in the male genital system of the schizopod My& jlexuosua. Journal of Cell Biology, 59, 284a. Reger, J. F., Dudkiewicz, A. B. and Florendo, N. (1970). The fine structure of spermatid-associated extracellulose tubules in the schizopod My& oculatu relicta. Journal of Ultrastructure Research, 30, 166-171. Retzius, G. (1909). I. Die Spermien der Schizopoden. In “Die Spermien der Crustacean”, pp. 23-29. Biologische Untersuchungen, new Series, 14. Reynolds, J. B. and De Graeve, G. M. (1972). Seasonal population characteristics of the opossum shrimp, Mysis relicta, in southeastern Lake Michigan, 1970-71. Proceedings of the Fvteenth Conjerence on Great Lakea Research, 117-131. Reys, J. P. (1960). Etude de la nourriture de quelques poissons demersaux du Golfe du Lion. Recueil des Travaux de la Station Marine d’Endoum, Facult6 des Sciences de Marseille, 33, 65-98. Rice, A. L. (1961). The response of certain mysids to changes in hydrostatic pressure. Journal of Experimental Biology, 38, 391-401. Rice, A. L. (1964). Observations on the effects of changes of hydrostatic pressure on the behaviour of some marine animals. Journal of the Marine Biological Association of the United Kingdom, 44, 163-176.

REFERENOES

307

Rice, A. L. (1967). The orientation of the pressure response of some marine Crustacea. I n “Symposium on Crustacea”, 3, pp. 11261131. Marine Biological Association of India. Richard, J. (1904). Campagne scientifique du yacht “Princesse-Alice” en 1903. Observations sur la sardine, sur le plankton, sur les c6tac68, sur des filets nouveaux, etc., etc. Bulletin du Mwke Ochnographique de Monaco, No. 11, 1-26.

Richards, R. C., Goldman, C. R., Frantz, T. C. and Wickwire, R. (1975). Where have all the Daphnia gone? The decline of a major cladoceran in Lake Tahoe, California-Nevada. Verhndlungen der Internationalen Vereinigung filr Theoretiache und Angewandte Limnologie, 19, 835-842. Richards, S. W. (1963). The demersal fish population of Long Island Sound. Bulletin ofthe Singham OceanographicCollection, Yale University, 18 (2), 1-101. Richards, S. W. and Riley, G. A. (1967). The benthic epifauna of Long Island Sound. Bulletin of the Bingham Oceanographic Collection, 19 (2), 89-129. Ricker, K. E. (1959). The origin of two glacial relict crustaceans in North America, as related to Pleistocene glaciation. Canadian Journal of Zoology, 37, 871-893.

Riedl, R. and Ozretic, B. (1969). Hydrobiology of marginal caves. Part I. General problems and introduction. Internationale Revue der Gesmten Hydrobiologie u. Hydrographie, 54,661-683. Ristich, S. S., Crandall, M. and Fortier, J. (1977). Benthic and epibenthic macroinvertebrates of the Hudson River. I. Distribution, natural history and community structure. Estuarine and Coa-~talMarine Science, 5 , 255-266. Robertson, A., Powers, C. F. and Anderson, R. F., (1968). Direct observations on My& relicta from a submarine. Limnology and Oceanography, 18, 700-702. Ross, S. T. (1977). Patterns of resource partitioning in searobins (Pisces: “riglidae). Copeia, 561-571. Ross, S. T. (1978). Trophic ontogeny of the leopard searobin, Prionotw, acitulw, (Pisces: Triglidae). Fishery Bulletin, National Oceanic Atmspheric Adminktration of the United States, 76, 225-234. Russell, F. S. (1925). The vertical distribution of marine macroplankton. An observation on diurnal changes. Journal of the Marine Biological Association of the United Kingdon, 11, 769-809. Russell, F. S. (1934). The zooplankton. 111. A comparison of the abundance of zooplankton in the Barrier Reef Lagoon with that of some regions in Northern European waters. Scientijic Repor& of the Great Barrier Reef Expedition, 2 , 176-201.

Rustad, D. (1930). Mysidacea. Scientiific Results of the Norwegian Antarctic Expeditions, 1927-29, No. 6, 1-28. Saether, 0. A. and McLean, M. P. (1972). A survey of the bottom fauna in Wood, Kalamalka and Skaha Lakes in the Okanagan Valley, British Columbia. Fkheries Research Board of Canada, Technical Report, No. 342, 1-27. Sage, L. E. and Herman, S. S. (1972). Zooplankton of the Sandy Hook Bay Area, N. J. Chesapeake Science, 13, 29-39. Sale, P. F., McWilliam, P. S. and Anderson, D. T. (1978). Faunal relationships among the near-reef zooplankton at three locations on Heron Reef, Great Barrier Reef, and seasonal changes in this fauna. Marine Biology, 49,133-145. Salonen, K., Sarvala, J., Hakala, I. and Viljanen, M.-L. (1976). The relation of energy and organic carbon in aquatic invertebrates. Limnology and Oceanography, 21, 726730.

308

TIIE BIOLOGY OF MYSIDS

Salvat, B. (1962). F a m e des sediments meubles intertidaux du Bassin d‘kcachon. Systematique et &ologie. Cahiers de Bwlogie Marine, 3, 219-244. Sando, H.(1964). Faunal list of the Zostera murina region at Kugurizaka coastal waters, Aomori Bay. Bullelin of the Biological Station of Asamuahi Tohoku University, 12, 27-35. Santa, N. and Bacescu, M. (1942). Recherche6 sur la vitamine C chez quelques crustaces marine. Bulletin de la Section Scienti&ue de I’Academie Roumaine, 24, 438-443. Sargent, J. R. (1976). The structure, metabolism and function of lipids in marine organisms. I n “Biochemical and Biophysical Perspectives in Marine Biology” (Eds D. C. Malins and J. R. Sargent), 3, pp. 150-212. Academic Press, New York and London. Sargent, J. R., Morris, R. J. and McIntosh, R. (1978). Biosynthesis of wax esters in oceanic crustaceans. Marine Biology, 46, 315-320. Siirkkii, J. (1972). The bottom macrofauna of the oligotrophic Lake Konnevesi, Finland. Annales Zoologici Fennici, 9, 141-146. Sarkka, J. (1976). Records of relict crustaceans in lakes drained by the River Kymijoki, Finland. Anna& Zoologici Fennici, 13, 44-47. Sam, G. 0. (1877). Nye bidrag ti1 Kundskaben om Middelhavets vets Invertebratfauna. I. Middelhavets Mysider. Archiv for Mathemtik og Naturvidenakab, 2, 10-119. Sm?, G. 0. (1885). Report on the Schizopoda collected by H.M.S. “Challenger” during the years 1873 to 1876. The Voyage of H.M.S. Challenger, 13, 1-228. Sars, G. 0. (1895). Crustacea Caspia. Account of the Mysidae in the collection of Dr 0. G r i m . Bulletin de 1’AcademieImperiale des Sciences de St. Petersbourg, 3, 433-458. Sars, G. 0. (1898). Isopoda. I n “An Account of the Crustaoea of Norway, 2, Bergen Museum. Sam, G. 0. (1907). Mysidae. Report of the Caspian Ezpdition, 1901, 1, 243-313. Saw, G. 0. (1916). On the juvenile state of Lophogaster typicw M. Sars. Archiv for Mathelnatik og Naturvidenskab, 34, 1-9. Sasaki, S. (1966a). Distribution of young striped bass, Roccua aaxatilis, in the Sacramento-San Joaquin Delta. California Fish and Game, 136,44-58. Sasaki, S. (1966b). Distribution of juvenile striped bass, Roccua saxatilds, in the Sacramento-San Joaquin Delta, California Fish ahd Game, 136, 59-07. Sasaki, S. ( 1 9 6 6 ~ )Distribution . and food habits of king salmon, Oncorhynchue tdmwytacha, and steelhead rainbow trout, Salrno gairdnerii, in the SacramentoSan Joaquin Delta. California Fish and Game, 136, 108-114. Saudray, Y. (1972). Consequencesdu cycle demue et d’intermue sur la morphologie fonctionnelle des pars molaris des Crustact58 Peracerid&. Revue dea Travaw, Institut des P&hes, Nantes, 36, 271-273. Savage, R. E. (1931). The relation between the feeding of the herring off the east coast of England and the plankton of the surrounding waters. Fishery Investigations, Ministry of Agriculture, Food and Fisheries, Ser. 2, 12 (3), 1-88. Savchuk, M. Ya. (1975). Feeding habits of sturgeon under present regime in the Sea of Azov. Trudg Vsesoyuznogo Nauchnoiasledovatel’skogo Imtituta Morskogo Rgbnogo Khozyaistva i OkeanograJii,59, 164-181. Savolainen, E. (1975). Distribution and food of Myoxocephalua quadricomis (L.) (Teleostei, Cottidae) in fresh waters of eastern Finland. Annalee Zoologki Pennici, 12, 271-274.

REFERENOES

309

Schaefer, R. H. (1970). Feeding habits of striped bass from the surf waters of Long Island. New York E’ish’and Game Journal, 17, 1-17. Schimmel, S. C:, Patrick, J. M. and Forester, J. (1977). Toxicity and bioconcentration of BHC and lindane in selected estuarine animals. Archives of Envirorvmental Contamination and Toxicology, 6 , 355-363. Schlkper, C. (1968). High pressure effects on marine invertebrates and fish. Marine Biology, 2, 5-12. Schneck, D., (1978). Comments on sampling tests made with Beyer’s epibenthic closing net. Meeresforschung, Reports on Marine Research, 26, 1114.

Schneider, J. C., Hooper, F. F. and Beeton, A. M. (1969). The distribution and abundance of benthic fauna in Saginaw Bay, Lake Huron. Proceedings of the Twelfth Conference on Great Lakes Research, 8-90. Schodduyn, M. (1926). Observations faites dans la Baie d ’ h b l e t e u s e (Pas de Calais). Bulletin de M w k e Ochnographique de Monaco, No. 482, 1-64. Schumacher, R. E. (1966). Successful introduction of Mysis relicta (Loven) into a Minnesota lake. Transactiom of the American Fisheries Society, 95, 216.

Scott, A. (1922). On the food of young plaice (Pleuronectea platessa). Journal of the Marine Biological Association of the United Kingdom, 12, 678-687. Scott, T. (1887). Natural history notes from Tarbert. Transactions of the Natural History Society of Glaagow, 1, 369-378. Scott, T. (1889). Notes on a few Crustacea and Mollusca new to the fauna of the Forth, with exhibition of specimens. Proceedings of the Royal Physical Society of Edinburgh, 10, 154-156. Scott, T. (1890). Additions t o the fauna of the Firth of Forth. Report of the Fishery Board for Scotland, 8 (3), 312-333. Scott, T. (1891a). The invertebrate fauna of the inland waters of Scotland. Report of the Fishery Board for Scotland, 9 (3), 269-296. Scott, T. (1891b). Additions to the fauna of the Firth of Forth. Report of the Fishery Board for Scotland, 9 (3), 300-310. Scott, T. (1894). Neomysis (Mysis) wulgaris J . V . Thompson in Barra, Outer Hebrides. Annals of Scottish Natural History, No. 12, 259. Scott, T. (1896a). Report on a collection of marine dredgings and other natural history materials made on the West Coast of Scotland by the late George Brook, F. L. S. Proceedings of the Royal Physical Society of Edinburgh, 13, 16s193.

Scott, T. (1896b). Erythrops serrata G. 0. Sars in the Firth of Clyde. Annals of Scottish Natural History, No. 19, 194. Scott, T. (1897). The marine fishes and invertebrates of Loch Fyne. Report of the Fishery Board for Scotland, 15 (3), 107-174. Scott, T. (1898). Some additions to the invertebrate fauna of Loch Fyne. Report of the Fishery Board for Scotland, 16 (3), 261-272. Scott, T. (1899). Report on the marine and freshwater Crustacea from FranzJosef Land, collected by Mr William S. Bruce of the Jackson-Harmsworth Expedition. Journal of the Linnean Society, Zoology, 27, 60-126. Scott, T. (19OOa). Notes on some gatherings of Crustacea collected for the most part on board the Fishery Steamer “Garland” and examined during the past year (1899). Report of the Fishery Board for Scotland, 18 (3), 382406.

Scott, T. (1900b). On the occurrence of Macrop&s slabberi (V.Ben.) in the Moray Firth. Annals of Scottish Natural History, No. 34, 126-127.

310

THE BIOLOGY OF MYSIDS

Scott, T. (190%). Notes on gatherings of Crustacea collected by the Fishery Steamer “Garland” and the steam trawlers “Star of Peace” and “Star of Hope” of Aberdeen, during the year 1901. Report of the Fishery Board for Scotland, 20 (3), 447-485. Scott, ! I (1902b). . Observations on the food of fishes. Report of the Fishery Board forScotland, 20 (1901, (3), 4 8 6 5 4 1 . Sedberry, G. R. and Musick, J. A. (1978). Feeding strategies of some demersal fishes of the continental slope and rise off the mid-Atlantic coa.st of the USA. Marine Biology, 44, 357-375. SegerstrAle, S. G. (1957). Baltic Sea. I n “Treatise on Marine Ecology and Palaeoecology”, (Ed. J. W. Hedgpeth), 1, pp. 751-800. Geological Society of America Memoir 67. Segerstrhle, S. G. (1962). The immigration and prehistory of the glacial relicts of Eurasia and North America. A survey and discussion of modern views. Internationale Revue der Geaamten Hydrobiologie u . Hydrographie, 47, 1-26, Seguin, G. (1968). Contribution a 1’6tude biochimique de Yraunus jfexuosua (Muller). Deep-sea Research, 15, 491-492. Sergeant, D. E. (1973). Feeding, growth and productivity of northwest Atlantic harp seals (Pagophilus groenbndicus). Journal of the Fisheries Research Board of Canada, 30, 17-29. Shelford, V. E. (1935). General survey of the communities, their extent and dynamics. The major communities. Ecological Monographs, 5 , 249-292. Shih, C.-T. and Laubitz, D. R. (1978). Zooplankton distribution in the eastern Beaufort Sea and the Northwest Passage. Astarte, Journal of Arctic Biology, 11, 45-54. Shotter, R. A. (1970). A new second intermediate host, Schistomysis spiritw (Sam), for the digenean, Podocotyl atomon (Rudolphi). Report of the Marine Biological Station, Port Erin, No. 82, 35-37. Shotter, R. A. (1972). Notes on helminth parasites of the whiting Odontagadus m. merlangus (L.)from the northern Irish Sea. Journal of Fish Biology, 4, 117-130. Shrivastava, H. (1974). Macrobenthos of Lake Huron. Fishery Research Board of Canada, Technical Report, No. 449, 1-45. Shurin, A. T. (1962). The distribution of bottom fauna in the eastern Baltic in 1960. Annales Biologiques, 17 (1960), 93-94. Shushkina, E. A. (1972). Intensity of production and utilization of assimilated food for growth of mysids from the Sea of Japan. Oceanology, 12, 275-285. Shushkina, E. A. (1973). Evaluation of the production of tropical zooplankton. I n “Life Activity of Pelagic Communities in the Ocean Tropics” (Ed. M. E. Vinogradov), pp. 172-183. Shushkina, E. A., Kuz’micheva, V. U. and Ostapenko, L. A. (1971). Energy equivalent of body mass, respiration and calorific value of mysids from the Sea of Japan. Oceanology, 12, 880-889. Shyamasundari, K. (1973). Mysidacea of Waltair coast. Rivista di Biologia dell’ Universita de Perugia, 66, 389-406. Siewing, R. (1953). Morphologische Untersuchungen an Tanaidaceen und Lophogastriden. Zeitschrift f u r Wissenschaftliche Zoologie, 157, 333-426. Siewing, R. (1956).Untersuchungen zur Morphologie der Malacostraca (Crustacea). Zoologische Jahrbucher, Anatomie und Ontogenie der Tiere, 7 5 , 39-176. Sikora, W. B., Heard, R. W. and Dahlberg, M. D. (1972). The occurrence and food habits of two species of hake, Urophycis regius and U . floridanus in Georgia estuaries. Transactions of the American FisheriesSociety, 101,513-826.

REFERENOES

311

Silas, E. G . and Pillai, P. P. (1975). Dynamics of zooplankton in a tropical estuary (Cochin Backwater), with a review on the plankton fauna of the environment. Bulletin of the Department of Marine Sciences, University of Cochin, 7, 329-355. Silva, V. M. A. P. da (1970a). Gaatrosaccw brasiliensis nova especie de Crustacea Mysidacea. A h da S'ociedade de Biologia do R i o de Janeiro, 13, 36-36. Silva, V. M. A. P. da (1970b). Metamysidopsk m a e n s i s nova espocie de Crustacea Mysidacea. Atas da Sociedu.de de Biologia do Rio de Janeiro, 13, 199-201. Silva, V. M. A. P. da (19'11e). Bowmanielkz (Bowmaniellu) atlantica nova nome para Umtrosaccw braailiensis (da Silva, 1970) (Crustacea, Mysidacea). Atm da Sociedade de Biologia do Rio de Janeiro, 14, 159. Silva, V. M. A. P. da (1971b). Bowmanielkz (Bownutniella) recifensis n.sp. Crustacea, Mysidacea. Atas de Sociedade de Biologia do Rio de Janeiro, 14, 185-187.

Silva, V. M. A. P. da (1972). Bowmaniella (Coifmaniella) inarticulatu sp.n. de Crustacea Mysidacea. Atm de Sociedade de Biologia do Rio de Janeiro, 15, 161-163.

Silva, V. M.A. P. da (1974). Siriella melloi, nova especie de Crustacea Mysidacea. Avuleo Departmento Zoologia Inetituto Biologia, Rio de Janeiro, 22, 1-4. Silva, V. M. A. P. da (1977). Mysidacea (Crustacea) da comissao nortelnordeste I . Dissertacao de Mestrado apre sentada a coordenacao do cur so de Posgraduacao em Zoologia da Rio de Janeiro, 1-93. Simmons, M. A. and Knight, A. W. (1975). Respiratory response of Neomysis intermedia (Crustacea : Mysidacea) to changes in salinity, temperature and season. Comparative Biochemistry and Phyeiology, 50, 181-193. Simmons, M. A., Sitts, R. M., Allen, J. T. and Knight, A. W. (1974). The nomenclature of mysids in the Sacramento-San Joaquin Delta Estuary. California Fish and Uame, 60, 23-25. Siudzinski, K. (1964). Macro- and ichthyoplankton of the southern Baltic in the period 1956-1959. A n n a k s Biologiquea, 19 (1962), 70-71. Sivertsen, E. (1938). Intersokelser over forholdet mellem spissog bredikodet a1 og deres naming. Report on the Norwegian F i s h e y and Marine Investigations, 5 (8), 1-22.

Skalkin, V. A. (1959). Food and food interrelationships of flounders of the Il'insk Bank. Investiya Tikhoolceanskogo nauchno-isseldovatel'skogo instituta gbnogo Khozyaistva i okeanografii, 47, 62-75. (Fishery Research Board of Canada Translation No. 789). Skjoldal, H. R. and Bamstedt, U. (1977). Ecobiochemical studies on the deepwater pelagic community of Korsfjorden, western Norway. Adenine nucleotides in zooplankton. Marine Biology, 42, 197-211. Smidt, E. L. B. (1951). Animal production in the Danish Waddensee. Meddekleer j r a Kommzissionen for Danmarks Fiakeri-og Havundersvgeleer, Serie : Fiekeri, 11 (6), 1-151.

Smith, S. I. (1879). The stalk-eyed crustaceans of the Atlantic coast of North America north of Cape Cod. Transactions of the Connecticut Academy of Arts and Sciences, 5, 27-138. Smith, W. E. (1970). Tolerance of Mysis relictu to thermal shock and light. Transactions of the American Fiaheries Society, 99, 418-422. Sparrow, R. A. H., Larkin, P. A. and Rutherglen, R. A. (1964). Successful introduction of My& relicta Loven into Kooteney Lake, British Columbia. Journal of the Fisheries Research Board of Canada, 21, 1325-1327.

312

THE BIOLOGY OF MYSIDS

Srinivasagam, R. T., R a F o n t , J. E. G., Moodie, C. F. and Raymont, J. K. B. (1971). Biochemical studies on marine zooplankton. X. The amino acid composition of Euphawia auperba, Meganyctiphanes norvegica and Neomyais integer. Journal of the Marine Biological Association of the United Kingdom, 51, 917-925. Stammer, H. J. (1933). Einige seltene oder neue Hohlentiere. Zoologischer Anzeiger, Supp. 6, 263-266. Stebbing, T. R. R. (1905). South African Crustacea. Part 111.Marine Investigations in South Africa, 4, 21-120. Stebbing, T. R. R. (1908). South African Crustacea. Part IV. Annah of the South African Muaeum, 6, 1-96. Stebbing, T. R. R. (1914). Stalk-eyed Crustacea Malacostraca of the Scottish National Antarctic Expedition. Transactions of the Royal Society of Edinburgh, 50, 253-307. Stebbing, T. R. R. (1917). South African Crustaceans. Part IX. Annals of the South African Museum, 17, 23-46. Stephensen, K. (1927). Papers from Dr Th. Mortensen’s Pacific Expedition 191P16. XL. Crustacea from the Auckland and Campbell Islands. Videmkabelige Meddeleher fra Dansk Naturhistoriak Forening i Kjobenhavn, 83, 289-390. Stephensen, K. (1933). The Scoresby Sound Committee’s 2nd East Greenland Expedition in 1932 to King Christian IX’s Land: Crustacea and Pycnogonida. Meddelelser om Grenland, 104 (15), 1-12. Stephensen, K. (1943). The zoology of East Greenland. Leptostraca, Mysidaoea, Cumacea, Tanaidacea, Isopoda and Euphausiacea. Meddelelser om Grenland, 121 (lo), 1-82. Steven, D. M. (1961). Shoaling behaviour in a mysid . Nature, London, 192, 280-281. Stevens, D. E. (1966a). Food habits of striped bass, Roccua sazatilia, in the Sacramento-San Joaquin Delta. California Fish and Game, 136, 68-96. Stevens, D. E. (1966b). Distribution and food habits of the American shad, Alosa sapidissima, in the Sacramento-San Joaquin Delta. Calqornia Fish and Game, 136, 97-107. Stickney, R. R . and Knowles, S. C. (1975). Summer zooplankton distribution in a Georgia estuary. Marine Biology, 33, 147-154. Stickney, R. R., Taylor, G. L. and White, D. B. (1975). Food habits of five species of young southeastern United States estuarine Soaenidae. Chesapeake Science, 16, 104-114. Stringer, G.E. (1967). Introduction of Mysis relicta Loven into Kalamalka and Pinaus Lakes, British Columbia. Journal of the Fisheries Research Board of Canada, 24, 463-465. Surtu, P. and Vives, F. (1957). Sobre la presensia de cfaatrosaccue sanctus (Van Beneden) en la desembocadura del rio Ebro. Inveetigacion Pmquera, 8, 121-126. Suomalainen, H. 0. T. (1954). The chromosome number of Neomysis vulgaris L. (Crustacea, Mysidae). Hereditas, 40, 3-4. Swain, W. R., Magnuson, R. W., Johnson, J . D., Olson, T. A. and Odlaug, T. 0. (1970). Vertical migration of zooplankton in western Lake Superior. Proceedings of the Thirteenth Conference on Great Lakes Research, 619-639. Swartz, R. C. (1972). Biological criteria of environmental change in Chesapeake Bay. Chesapeake Science, 13 (Supp.), 517-541.

REFERENOES

313

Taberly, G. (1954a). Etude morphologique d‘un Dajidae peu connu: Prodajus Zobiancoi Bonnier (Crust. Isop. ,Epictiride). I. L’epicaridium de P. Zobiancoi et remarques g6n6rales sur l’epicaridium des Dajidae. Bulletin de Musde Ochnographique de Monaco, No. 1045, 1-9. Taberly, G. (1954b). Etude morphologique d‘un Dajidae peu connu: Prodajus lobiancoi Bonnier (Crust. Isop. Epicaride). 11.Le cryptoniscium de P. lobiancoi et sa mue formes connues de cryptoniscium de Dajidae. Bulletin de M w d e Ocdanographique de Monaco, No. 1049, 1-15. Taniguchi, A. (1969). Mysidacea and Euphausiacea collected in the south-east of Hokkaido, Japan. Bulletin of the Faculty of Fisheries, Hokkaido University, 20, 43-59. Taniguchi, A. (1976). Mysids and euphausiids in the eastern Indian Ocean with particular reference to invasion of species from the Banda Sea. Journal of the Marine Biological Association of India, 6, 349-357. Tattersall, 0. S. (1955). Mysidacea. “Discovery” Reports, 28, 1-190. Tattersall, 0. S. (1957). Report on a small collection of Mysidacea from the Sierra Leone Estuary together with a survey of the genus Rhopalophthalmus Illig and a description of a new species of Tenagomysis from Lagos, Nigeria. Proceedings of the Zoological Society of London, 129, 81-128. Tattersall, 0. S. (1958). Further notes on Mysidacea from South African waters. Transactions of the Royal Society of South Africa, 35, 373-385. Tattersall, 0. S. (19604. Report on a small collection of Mysidacea from Singapore waters. Proceedings of the Zoological Society of London, 135, 165-181. Tattersall, 0. S. (1960b). Notes on mysidacean crustaceans of the genus Lophogaster in the U.S. National Museum. proceedings of the United States National Museum, 112, 527-547. Tattersall, 0. S. (1961a). Report on some Mysidacea from the deeper waters of the Ross Sea. Proceedings of the Zoological Society of London, 137,553-571. Tattersall, 0.S. (1961b). Mysidacea from the coast of tropical West Africa. Atlamtide Report, 6, 143-159. Tattersall, 0. S. (1962). Report on a collection of Mysidacea from South Africa off-shore and coastal waters (1957-1959) and from Zanzibar. Proceedings of the Zoological Society of London, 139, 221-247. Tattersall, 0. S. (1965a). Report on a small collection of Mysidacea from the northern region of the Malacca Strait. Journal of Zoology, 147, 75-98. Tattersall, 0.S. (1965b). The fauna of the Ross Sea. Part 4. Mysidacea. B u l k t i n of the N e w Zealand Department of Scientific and Industrial Research, 167, 1-35. Tattersall, 0. S. (1967).A survey of the genus Heteromysis (Crustacea: Mysidacea) with description of five new species from tropical coastal waters of the Pacific and Indian Oceans, with a key for the identification of the known species of the genus. Transactions of the Zoological Society of London, 31, 157-193. Tattersall, 0.S. (1968). A new species of Petalophthulmus (Mysidacea) based on specimens from off Puerto Rico, hitherto referred to as Petalophthalmus oculatus. Journal of Zoology, 155, 271-282. Tattersall, 0.S. (1969).A synopsis of the genus Mysidopsis (Mysidacea, Crustacea) with a key for the identification of its known species and description of two new species from South African waters. Journal of Zoology, 158, 63-79. Tattersall, W. M. (1908). The fauna of brackish water ponds a t Port Canning, Lower Bengal. Part XI. Two new Mysidae from brackish water in the Ganges Delta. Record of the Indian Museum, 2, 233-239.

314

THE BIOLOGY OF MYSIDS

Tattersall, W. M. (1919). Notes on Lancashire and Cheshire specimens. 1. Crustacea from pools near Leasow Lighthouse on the Wirral Peninsula. Lancashire and Cheshire Naturalist, 12, 20-26. Tattersall, W. M. (1921). Mysidacea, Tanaidacea and Isopoda. Pt. 7. Memoirs of the Asiatic Society of Bengal, 6, 403-433. Tattersall, W. M. (1922). Indian Mysidacea. Records of the Indian Muaeum, 24, 445-504. Tattersall, W . M. (1938). The seasonal occurrence of mysids off Plymouth. Journal of the Marine Biological Association of the United Kingdom, 23, 43-56. Tattersall, W . M. (1951). A review of the Mysidacea of the United States National Museum. Bulletin of the United States National Museum, 201, 1-292. Tattersall, W. M. and Tattersall, 0. S. (1951). “The British Mysidacea”, 460 pp. Ray Society Monograph. Tchindonova, J. G. (1959). Feeding of some groups of macroplankton in the northwestern Pacific. Trudg Instituta Okeanologii, Akademiya Nauk SSSR, 30, 166-189. Teraguchi, M., Wissing, T. E. and Hasler, A. D. (1972). Change in caloric content of adult males of Mysis relicta (Loven) during a die1 migratory cycle. American Midland Naturalist, 88, 235-239. Teraguchi, M., Hasler, A. D. and Beeton, A. M. (1975). Seasonal changes in the response of Mysis relicta Loven to illumination. Verhandlungen der Internationalen Vereinigung f ilr theoretische und angewandte Limndogie, 19, 2989-3000. Teshima, S. and Kanazawa, A. (1971). Sterol composition of marine crustaceans. Bulletin of the Japanese Society of Scientific Fisheries, 37, 63-67. Tett, P. B. and Kelly, M. G. (1973). Marine bioluminescence. Oceanography and Marine Biolagy, a n Annual Review, 11, 89-173. Thomas, J. L. (1967). The diet of juvenile and adult striped bass, Roccwr suxatil49, in the Sacramento-San Joaquin River System. California Fish and Game, 53, 49-62. Thommes, M. M., Lucas, H. F. and Edgington, D. N. (1972). Mercury concentrations in fish taken from offshore areas of the Great Lakes. Proceedings of the Fifteenth Conference on Great Lakes Research, 192-197. Todd, R. A. (1914). Report on the food of the plaice. Fishery Inweatigationa, Ministry of Agriculture, Food and Fisheries, Ser. 11, 2, 1-31. Tracy, S. F. and Vallentyne, J. R. (1969). Fungal decomposition and amino acid analysis of Mysis relicta Loven. Limnology and Oceanography, 11, 352-356. Tskhomelidze, 0. I. and Sergeeva, Zh. N. (1966). Acclimatization of food invertebrates in the waters of Georgia, USSR. Voprosg Ikhtiologii, 6, 752-753. Tundisi, J. and Krishnaswamy, S. (1967). Total phosphorus content of Neomyab integer. Boletim do Institute Oceanogra$co, Sao Paulo, 16, 99-100. Turner, J. L. (1966a). Distribution and food habits of ictalurid fishes in the Sacramento-Sen Joaquin Delta. California Fish and Game, 136, 130-143. Turner, J. L. (196613). Distribution and food habits of centrarchid fishes in the Sacramento-San Joaquin Delta. California Fish and G u m , 136, 144-153. Turner, J. L. and Heubach, W. (1966). Ecological studies of the Sacramento-San Joaquin Estuary. Distribution and concentration of Neomysia awatschewis in the Sacramento-SanJoaquin Delta. California Fish and Game, 133, 105-112. Tyler, A. V. (1972). Food resource division among northern marine, demersd fish. Journal of the Fisheries Research Board of Canada, 29, 997-1003.

REFERENCES

316

Tyler, A. V. (1973). Caloric values of some North Atlantic invertebrates. Marine Biology, 19, 258-261. ’ Uzars, D. (1969). Feeding conditions of Baltic cod in 1967 and 1968. Anruzh Biologiques, 25 (1968), 121-122. Uzars, D. (1972). Feeding and relative liver weight of Baltic cod in the winterspring period of 1970. Annalm Biologiqum 27 (1970), 92-93. Vader, W. (1973). A bibliography of the Ellobiopsidae, 1959-1971, with a list of Thabsomyces species and their hosts. Sarsia, 5 2 , 175-180. Valkanov, A. (1957). Katalog unserer Schwarzmeerfauna. Arbeiten a m der Biologischen Meeresstation in Varna (Bulgarien), 19, 1-62. Vannini, E. (1930). Nota sulk forme larvali di Leptomysis lingvura del Golfo di Napoli. Pubblicazioni della Stazione Zoologica di Napoli, 10, 285-296. Vinogradov, G. A. (1976). Osmotic regulation of some glacial-relict crustaceans in connection with their ecology and origin. I n “Explorations of the Fauna of the Seas XVII (XXV). Salinity Adaptations of the Aquatic Animals”, pp. 167-210. Zoological Institute, Academy of Sciences of the USSR. Vinogradov, M. E. (1962). Feeding of the deep-sea zooplankton. Rapport et proces-verbaux,dm rdunions. Conaeil permnent inte7national pour 1 ’exploration de la mer, 153, 114-119. Vinogradov, M. E. (1970a). Vertical Dwtribution of the Oceanic Zooplankton. Israel Program for Scientific Translations, Jerusalem, 1-339. Vinogradov, M. E. (1970b). The vertical distribution of zooplankton in the Kurile-Kamchatka region of the Pacific Ocean (based on data from the 39th cruise of the R/V “Vityaz”). Trudg Imtitutu Okeanologii, Akademiya Nauk, SSSR, 86, 99-116. Vinogradov, M. E. and Parin, N. V. (1973). Some features of the vertical ’ distribution of macroplankton in the tropical Pacific. Okeanologiia, 13, 137-148. Vinogradov, M. E., Bordovskiy, 0. K. and Akhmet’yeva, Ye. A. (1970). Biochemistry of oceanic plankton and chemical composition of plankton from different depths of the northwestern Pacific. Oceanology, 10, 692-698. Vinogradov, Z. A. and Kandiuk, R. P. (1967). Sterols, provitamins D and cholesterol in marine organisms. I n “Biochemistry of Marine Organisms”, pp. 61-69. Ukrainian Academy of Sciences, USSR. Vinogradova, Z. A. and Petkevich, T. A. (1967). Chemical Composition of the plankton of the Black, Azov and Caspian Seas. I n “Biochemistry of Marine Organisms”, pp. 70-82. Ukrainian Academy of Sciences, USSR. Vives, F. (1965). Rapports entre hydrographie et zooplankton dans m e region neritique de la MBditerranBe occidentale. Rapport et prom-verbaux dea riuniona. Cornmiasion intemationale pour l’exploration scientifiue de la mer MeZiterranke, 18, 383-389. Vives, F. (1968). Sur les malacostraces planctoniques des mere Tyrrhenienne et Catalane. Rapport et proces-verbaux des riuniona. Cornmiasion internutionale pour l’exploration scientihue de la mer MdditerranBe, 19, 459-461. Vlasblom, A. G. and Elgershuizen, J. H. B. W. (1977). Survival and oxygen consumption of Praunwr jlexmswr and Neomysis integer, and embryonic development of the latter species, in different temperature and chlorinity combinations. Netherlands Journal of Sea Research, 11, 305-315. Vlymen, W. J. (1970). Energy expenditure of swimming copepods. Limnology and Oceanography, 15, 348-356.

316

THE BIOLOGY OF MYSIDS

Vogt, W. (1932). Uber die Morphologie und Histologie der Antennendruse und der thoracalen Arthrocytenorgane der Mysideen. Zeitmhrift f u r Morphologie und Okologie der Tiere, 24, 288-318. Vogt, W. (1933). Uber die Antennendrusevon My& relicta. Zoologische Jahrbwher, Anatomie und. Ontogenie der Tiere, 56, 373-386. Vorstman, A. G. (1951). A year’s investigation on the life cycle of Neomysia vulgar& Thompson. Verhandlungen der Internationalen Vereinigung f u r theoretische und angewandte Limnologie, 11, 437-445. Vos, A. P. C. (1941). Zoologische resultaten van een Tocht rond het Ijsselmeer van 5-8 Juli 1937. De Biologk van de Zuidenee Tijdena h r Drooglegging, 5, 37-50.

Wagner, J. (1896). Einige Beobachtungen iiber die embryonale Entwicklung von Nwrnysia vulgaria var. baltica Czerniavsky. Travaux de la Soci6t6 I r n p W e des Naturalistes de St. Petersbourg, Section de Zoologie et Physiologie, 26, 177-221.

Walker, A. 0. (1886). Report on the Podophthalmata of the L.M.B.C. district. Report of the Liverpool Marine Biological Committee, 1, 221-226. Walker, A. 0. (1888a). Report on the Crustacea of Liverpool Bay 1886-1887. Proceedings of the Liverpool Biological Society, 2, 171-181. Walker, A. 0. (188813). Report on some h s t a c e a from the Canary Islands collected by Messrs I. C. Thompson and W. S. McMillen. Proceedings of the Liverpool Biological Society, 2, 130-132. Walker, A. 0. (189%). Malacostraca from Puget Sound, N.W. America. Anand Magazine of Natural History, Ser. 7, 2, 275-276. Walker, A. 0. (189813). Malacostraca from the west coast of Ireland. Proceedings and Transactiona of the Liverpool Biological Society, 12, 159-172. Walton, J. (1922). A Spitsbergen salt marsh: with observations on the ecological phenomena attendant on the emergence of land from the sea. J w m d of Ecology, 10, 109-121. Warwick, R. M. and Davies, J. R. (1977). The distribution of sublittoral macrofauna communities in the Bristol Channel in relation to the substrate. Estuarine and Coastal Marine Science, 5, 267-288. Waterman, T. H. (1955). Polarized light and animal navigation. Scientij’k American, 193, 8&94. Waterman, T. H., Nunnemacher, R. F., Chace, F. A. and Clarke, G. L. (1939). Diurnal vertical migrations of deep-water plankton. Biological Bulletin, Marine Biological Laboratory, Wood’s Hole, Mass., 76, 256-279. Watson, N. H. (1974). Zooplankton of the St Lawrence Great Lakes-species composition, distribution and abundance. Journal of the Fisheries Reaearch Board of Canada, 31, 783-794. Weish, P. and Tiirkay, M. (1975). L i m n m y & benedeni in Osterreich mit Betrachtungen zur Besiedlungsgeschichte (Crustacea: Mysidacea). Archiv f fir Hydrobwlogie, Supp. 44, 48&491. Wells, L. (1960). Seasonal abundance and vertical movements of planktonic Crustacea in Lake Michigan. Fishery Bulletin, Fish and Wildlye Service, United States Department of the Interior, 60, 339-369. Wells, L. and Beeton, A. M. (1964). Food of the bloater, Coregmwr h q i , in Lake Michigan. Tramactions of the American Fisheries Society, 92, 246255.

Wenner, A. M. (1972). Sex ratio as a function of size in marine Crustacea. American Naturalist, 106, 321-350.

REFERENUES

317

Werner, A. M., Fusaro, C. apd Oaten, A. (1974). Size a t onset of maturity and growth rate in crustacean populations. Canadian Journal of Zoology, 52, 1095-1106.

Westin, L. (1970). The food ecology and the annual food cycle in the Baltic population of fourhorn sculpin, Myoxocephalus quadricornis (L.) Pisces. Report of the Institute of Freshwater Research, Drottningholm, 50, 168-210. Whiteley, G. C. (1948). The distribution of larger planktonic Crustacea on Georges Bank. Ecological Monogrqhs, 18, 233-264. Whitfield, A.K.andBlabor, S.J.M. (1978). Food andfeeding ecology ofpiscivorous fishes at Lake St Lucia, Zululand. Journal of Fish Biology, 13, 675-691. Wiborg, K. F. (1971). Investigations on euphausiids in some fjords on the west coast of Norway in 1966-1969. Fiskeridirektoratets Skrifter, Serie Havundersekelser, 16, 10-35. Wickstead, J. H. (1953). A new apparatus for the collection of bottom plankton. Journal of the Marine BiologicalAssociationof the United Kingdom, 32,347-355. Wickstead, J . H. (1961). A quantitative and qualitative study of some Indo-WestPacific plankton. Fishery Publications, Colonial Oflce, No. 16, 1-200. Wiens, A. P., Rosenberg, D. M. and Snow, N. B. (1975). Species list of aquatic plants and animals collected from the Mackenzie and Porcupine River watersheds from 1971 to 1973. Fisheries and Marine Service Research and Development Directorate, Department of the Environment, Canada, Technical Report No. 557, 1-39. Wigley, R. L. (1963). Occurrence of PraunwrJlexuosus (0.F. Muller) (Mysidacea) in New England waters. Cruataceana, 6, 158. Wigley, R. L. and Burns, B. R . (1971). Distribution and biology of mysids (Crustacea, Mysidacea) from the Atlantic coast of the United States in the NMFS Wood’s Hole Collection. Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 69, 717-746. Wiktor, K. (1961). Observations on the biology of N e o m y k vulgaris (Thompson) in Zalew Szczecinski (Stettiner Haff) and Zatoka Pomorska (Pomeranian Bay). Przeglud Zoologiczny, 5, 36-42. Williams, A. B. (1972). A ten-year study of meroplankton in North Carolina estuaries: mysid shrimps. Chesapeake Science, 13, 254-262. Williams, A. B., Bowman, T. E. and Damkaer, D. M. (1974). Distribution, variation and supplemental description of the opossum shrimp, N m y s i s americana (Crustacea : Mysidacea). Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 72, 835-842. Wilson, J. B. and Roff, J. C. (1973). Seasonal vertical distributions and diurnal migration patterns of Lake Ontario crustacean zooplankton. Proceedings of the Sixteenth Conference on Great Lakes Research, 190-203. Win, K . (1977). A species list of the zooplankton from the coastal waters of Burma. I n “Proceedings of the Symposium on Warm Water Zooplankton”, pp. 93-99. Special Publication of National Institute of Oceanography, Goa, India. Wing, B. L. (1965). A Thalaeaomyces (Ellobiopsidae) infesting Acanthomysia pseudomacropis and Neomysis kdiakensis (Mysidacea)in southwestern Alaska. American Society of Limnology and Oceanography, 1, 295. Wing, B. L. (1975). New records of Ellobiopsidae (Protista (incertae s e d i s ) ) from the North Pacific with a description of Thalmsmyces albatrossi n.sp., a parasite of the mysid Stylmysis major. Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 73, 169-185.

318

THE BIOLOGY OF MYSIDS

Wing, B. L. and Barr, N. (1977)..Midwater invertebrates from the southeastern Chukchi Sea: species and abundance in catches incidental to midwater trawling survey of fishes, September-October, 1970. National Oceanic Atmospheric Administration of the United States, Technical Report NMFS SSRF-710, 1-43. Wissing, T. E., Darnell, R. M., Ibrahim, M. A. and Berner, L. (1973). Caloric values of marine animals from the Gulf of Mexico. Contributions in Marine Science, 17, 1-17. Wittmann, K. J. (1977). Modification of association and swarming in north Adriatic Mysidacea in relation to habitat and interacting species. I n “Biology of Benthic Organisms” (Eds B. F. Keegan, P. 0. Ceidigh and P. J. S: Boaden), pp. 605-612. Pergamon Press, Oxford. Wittmann, K. J. (1978). Adoption, replacement and identification of young in marine Mysidacea (Crustacea). Journal of Experimental Marine Biology and Ecology, 32, 259-274. Wooldridge, T. (1977). The zooplankton of Mgazana, a mangrove estuary in Transkei, southern Africa. Zoologica Africana, 12, 307-321. Woynarovich, E. (1954). Vorkommen der Limnomysis benedeni Czern. im ungarischen Donauabschnitt . Acta Zoologica Academiae Scientarum Hungaricue, 1, 177-185. Yaguchi, E. M., Waller, B. J. and Marshall, J. S. (1974). Plutonium distribution in Lake Michigan biota. Proceedings of the Seventeenth Conference on Qreat Lakes Research, 150-157. Yamada, M. (1961). Studies on the lipid of plankton. I. The lipid of Neomysis nakazawai Ii. Journal of the Japan Oil Chemist’s Society, 10,23G239. Yamada, M. (1964). The lipid of plankton. Bulletin of the Japanese Society of Scientific Fisheries, 30, 673-681. Yamamoto, G. (1977). Successional processes observed in the freshening brackish lakes of Shimokita Peninsula. Journal of the Faculty of Marine Science and Technology, Tokai University, No. 10, 119-128. Zatkutskiy, V. P. (1965). The concentration of various benthic and demersal organisms in the near-surface layer of the Black and Azov Seas. Oceanology, 5, 83-85. Zatkutskiy, V. P. (1970). Some biological features of mysids of the hyponeuston in the Black Sea and Sea of Azov. Gidrobiologicheskii Zhurnal, 6 , 17-22. Zelickman, E. A. (1974). Group orientation in Neomysis mirabilk (Mysidacea: Crustacea). Marine Biology, 24, 251-258. Zelickman, E. A. and Golovkin, A. N. (1972). Composition, structure and productivity of neritic plankton communities near the bird colonies of the northern shores of Novaya Zemlya. Marine Biology, 17,265-274. Zenkevich, L. A. (1957). Caspian and Aral Seas. I n “Treatise on Marine Ecology and Palaeoecology” (Ed. J. W. Hedgpeth), 1, pp. 891-916. Geological Society of America, Memoir 67. Zenkevich, L. A. and Birstein, J. A. (1960). On the problem of the antiquity of the deep-sea fauna. Deep-sea Research, 7 , 10-23. Zharkova, I. S. (1970). Reduction of the organs of vision in deep-sea mysids. Zoologicheskii Zhurnal, 49, 685-693. Zhuravel, P. A. (1959). On the biology and ecology of mysids experimentally introduced into water reservoirs and other bodies of water in the Ukraine in order to enrich the feeds provision of fishes. Zoologicheskii Z h u m l , 38, 987-990.

REFERENCES

319

Zhuravel, P. A. (1960a). On new species in the fauna of the Bakhchisarai Water Reservoir in the Crimea.. Zoologicheekii Z h u m l , 39,458459. Zhuravel, P. A. (1960b), The mysid Hemimy& a n m l a Sars (Crustacea Malacostraca) in the Dnieper water reservoir and its feeding value for fishes. Zoologicheskii Zhurnal, 39, 1571-1573. Zhuravel, P. A. (1972). The Caspian estuarine mysid Mesomysk kowalewskyi Czern. in the Baltic Sea. Wrobiologicheskii Zhurnal, 8, 82-83. Zixnmer, C. (1912). Sudwestafrikanische Schizopoden. Jenaische Denkschriften, 17, 1-11.

Zimmer, C. (1918). Neue und wenig bekannte Mysidaceen des Berliner ZooIogischen Museums. Mitteilungen aus dem Zoologkchen M w e u m in Berlin, 9, 13-26.

Zmudzinski, L. (1962). The mysid P r a u n w i n e m i s (Rathke) in Puck Bay. Annales Biologiques, 17 (1960), 91. Zyblut, E. R. (1970). Long-term changes in the limnology and macrozooplankton of a large British Columbia lake. Journal of the Fisheries Research Board of Canada, 27, 1239-1250.

APPENDIX I

TAXONOMIC LIST OF MYSIDACEA The following is a taxonomic list of the Order of Mysidacea containing the integrated corrections and additions to the World List of Mauchline and Murano (1977). The genera are listed alphabetically under Families, Sub-Families or Tribes. The species in turn are also listed alphabetically for easy reference. Sub-order LOPHOGASTRIDA Family LOPHOGASTRIDAE Genus Ceratolepis G. 0. Sars, 1883 C. hamata G. 0. Sam, 1883 Genus Chabraspidum W.-Suhm, 1895 C. alatum ( W . - S u b , 1876) Genus Pseudochalaraspidum Birstein and Tchindonova, 1962 P. hanseni Birstein and Tchindonova, 1962 Genus Gnathophausia W.-Suhm, 1875 G. afinis G. 0. Sam, 1883 G. elegans G. 0. Sars, 1883 G. gigas W.-Suhm, 1875 G. gracilis W.-Suhm, 1875 G. ingens (Dohm, 1870) G. longispina G. 0. Sars, 1883 G. scapularis Ortmann, 1906 G. zoea W.-Suhm, 1875 Genus Lophogaster M. Sars, 1857 L. afi& Colosi, 1930 L. americanus W. Ta%tersall,1951 L. challengeri Fage, 1940 L. erythraeus Colosi, 1930 L. hawaiensis Fage, 1940 L. intermedius Hansen, 1910 L. japonicus W. Tattersall, 1951 L. longirostris Faxon, 1896 L. multispinosus Fage, 1940 L. pacijicus Fage, 1940

APPENDIX I

L. rotundatus Illig, 1930 L. schmidti Page, 1940 L. spinosus Ortmann, 1906 L. subglaber Hansen, 1927 L. typicus M. Sars, 1857 Genus Paralophogaster Hansen, 1910 P. atlanticus W. Tattersall, 1937 P. glaber Hansen, 1910 P.indicus Pillai, 1973 P.intermedius Coifmann, 1937 P. macrops Colosi, 1934 P. microps Colosi, 1930 P . sanzoi Colosi, 1930 Family EUCOPIIDAE Genus Eucopia Dana, 1852 E . australis Dana, 1852 E . grimaldii Nouvel, 1942 E . linguicauda 0. Tattersall, 1955 E . sculpticauda Faxon, 1893 E . unguiculata (W.-Suhm, 1875) Sub-order MYSIDA Family PETALOPHTHALMIDAE Genus Ceratomysis Faxon, 1893 C. egregia Hansen, 1910 C. ericula Ledoyer, 1977 C. spinosa Faxon, 1893 Genus Hansenomysis Stebbing, 1893 H . angusticauda 0. Tattersall, 1961 H . antarctica Holt and Tattersall, 1906 H . armata Birstein and Tchindonova, 1958 H . birsteini Bacescu, 1971 H . chini Bacescu, 1971 H . falklandica 0. Tattersall, 1955 H . fyllae (Hansen, 1887) H . lucifugus (Faxon, 1893) H . menziesi Bacescu, 1971 H . peruvianus Bacescu, 1971 H . rostrata Birstein and Tchindonova, 1970 H . spenceri Bacescu, 1971 H . tattersallae Bacescu, 1971 H. tropicalis Bacescu, 1967 H . violacea (Birstein and Tchindonova, 1958)

321

322

THE BIOLOGY OF MYSIDS

Genus Petalophthalmus W.-Suhm, 1875 P. armige? W.-Suhm, 1875 P . caribbeanus 0. Tahtersall, 1968 P. oculatus Illig, 1906 Family MYSIDAE Sub-family BOREOMYSINAE Genus Boreomysis G. 0. Sars, 1969 B. acuminata 0. Tattersall, 1955 B. arctica (Kr~yer,1861) B. atlantica Nouvel, 1942 B. bispinosa 0. Tattersall, 1955 B. brucei W. Tattersall, 1913 B. caeca Birstein and Tchindonova, 1958 B . californica Ortmann, 1894 B. cheZata Birstein and Tchindonova, 1958 B. curtirostris Birstein and Tchindonova, 1958 B. dubia Coifmann, 1936 B. fragilis Hansen, 1912 B . hanseni Holmquist, 1956 B. illigi 0. Tattersall, 1955 B. incisa Nouvel, 1942 B. inermis ( W . - S u b , 1874) B . insoZita 0. Tattersall, 1955 B. intermedia Ii, 1964 B. jacobi Holmquist, 1956 B. kistnae Pillai, 1973 B. latipes Birstein and Tchindonova, 1958 B . longispinu Birstein and Tchindonova, 1958 B. macrophthalma Birstein and Tchindonova, 1958 B. megalops G. 0. Sars, 1872 B. microps G. 0. Sars, 1883 B. nobilis G. 0. Sars, 1885 B. obtusata G. 0. Sars, 1883 B. plebeja Hansen, 1910 B. rostrata Illig, 1906 B. semicoeca Hansen, 1905 B. sibogae Hansen, 1910 B. sphaerops Ii, 1964 B. tanakai Ii, 1964 B. tatters& 0. Tattersall, 1955 B. tridens G . 0. Sam, 1870 B. vanhoefleni Zimmer, 1914

APPENDIX I

B. verrucosa W. Tattersall, 1939 Sub-family SIRIELLINAE Genus Hemisiriella Hansen, 1910 H . abbreviata Hansen, 1912 H . gardineri W. Tattersall, 1912 H . parva Hansen, 1910 H . pulchra Hansen, 1910 Genus Siriella Dana, 1850 S. adriatica Hoenigman, 1960 S. aequiremis Hansen, 1910 S. afinis Hansen, 1910 S. anomla Hansen, 1910 S. armata (M.-Edwards, 1837) S. australis W. Tattersall, 1927 S. brevicaudata Paulson, 1875 S. brevirostris Nouvel, 1944 S. castellabatensis Ariani and Spagnuola, 1976 S. chierchiae Coifmann, 1937 S. clausii G. 0. Sars, 1877 S. conformalis Hansen, 1910 S. dayi 0. Tattersall, 1952 S. denticulata (Thomson, 1880) S. distinguenda Hansen, 1910 S. dollfusi Nouvel, 1944 S. dubia Hansen, 1910 S. gibba Nouvel, 1944 S. gracilis Dana, 1852 S. halei W. Tattersall, 1927 S. hanseni W. Tattersall, 1922 S. inornata Hansen, 1910 S. jaltensis Czerniavsky, 1868 S. japonica Ii, 1964 S. jonesi Pillai, 1964 S. lingvura Ii, 1964 S. longidmtyla W. Tattersall, 1940 S. longipes Nakazawa, 1910 S. media Hansen, 1910 S. melloi Silva, 1974 S. mexicana Brattegard, 1970 S. nodosa Hansen, 1910 S. norvegica G. 0. Sars, 1869 S. okadai Ii, 1964

323

324

TEE BIOLOGY OF MYSIDS

S. paci$m Holmes, 1900 S. panamemis W. Tattersall, 1951 S. paulsoni Kossman, 1877 S. plumicauda Hansen, 1910 S . podoensis 0. Tatbersall, 1962 S. quadrispinosa Hansen, 1910 S. quilonensis Pillai, 1961 S. robusta Pillai, 1964 S. roosevelti W. Tattersall, 1941 S. serrata Hansen, 1910 S. sinensis Ii, 1964 S. singularis Nouvel, 1957 S. tadjourensis Nouvel, 1944 S. thompsonii (M.-Edwards, 1837) S. trispina Ii, 1964 S. vincenti W. Tattersall, 1927 S. vulgaris Hansen, 1910 S. wadai Ii, 1964 S. watasei Nakazawa, 1910 S . wolfJi 0. Tattersall, 1961 Sub-family RHOPALOPHTHALMINAE Genus Rhopalophthulmus Illig, 1906 R. africana 0. Tattersall, 1957 R. brisbanensis Hodge, 1963 R. chilkensis 0. Tattersall, 1957 R. dakini 0. Tattersall, 1957 R. egregius Hansen, 1910 R. jlagellipes Illig, 1906 R. indicus Pillai, 1961 R. kempi 0. Tattersall, 1957 R. longicauda 0. Tattersall, 1957 R. longipes Ii, 1964 R. macropsis Pillai, 1964 R. mediterraneus Nouvel, 1960 R. orientalis 0. Tattersall, 1957 R. tattersazlae Pillai, 1961 R. terranataEs 0. Tattersall, 1957 Sub-family GASTROSACCINAE Genus Anchialina Norman and Scott, 1906 A . agilis (G. 0. Sam, 1877) A. dantani Nouvel, 1944 A. dentata Pillai, 1964

AF'PENDIX I

A. Jlemingi W . Tattersall, 1943 A. grossa Handen, 1910 A . latifrons Nouvel, 1971 A. madagmcariensis Nouvel, 1969 A . media Ii, 1964 A. obtusifrons Hansen, 1912 A. oculata Hoenigman, 1960 A. penicillata Zimmer, 1915 A . sanzoi Coifinann, 1937 A . truncata (G. 0. Sam, 1883) A . typica (Kmyer, 1861) A. zimmeri W . Tattersall, 1951 Genus Archaeomysis Czerniavsky, 1882 A. grebnitzkii Czerniavsky, 1882 A. kokuboi Ii, 1964 Genus Bowmaniella Bacescu, 1968 B. atlantica Silva, 1971 B. bacescui Brattegard, 1970 B. banneri Bacescu, 1968 B. brmiliensis Bacescu, 1968 B. dissimilis (Coifmann, 1937) B. Jloridana Holmquist, 1975 B. inarticulata Silva, 1972 B. johnsoni (W. Tattersall, 1937) B. merjonesi Bacescu, 1968 B. mexicana (W. Tattersall, 1951) B. parageia Brattegard, 1970 B. portoricensis Bacescu, 1968 B. recifensis Silva, 1971 B. sewelli Brattegard, 1970 Genus Gastrosaccus Norman, 1868 G. australis W. Tattersall, 1923 G. bengalensis Hansen, 19 10 G. bravi$ssura 0. Tattersall, 1952 G. dalcini W . Tattersall, 1940 G. dunckeri Zimer, 1915 G. elegans 0. Tattersall, 1960 G. erythraeus (Kossman, 1877) G. formosensis Ii, 1964 G. gordonae 0. Tattersall, 1952 G. hibii Ii, 1964 G. indicw Hansen, 1910

325

326

THE BIOLOGY OF MYSIDS

G . kempi W. Tattersall, 1922 G. lobatus Nouvel’, 1951 G. magnilobatus Bacescu and Schiecke, 1974 G. mediterruneus Bacescu, 1970 G. msangii Bacescu, 1975 G. muticus W. Tattersall, 1915 G. normani G. 0. Sars, 1877 G. ohhimai Ii, 1964 G. olivae Bacescu, 1970 G. pmi$cus Hansen, 19 12 G. parvus Hansen, 1910 G. pelagicus Ii, 1964 G. psammodytes 0. Tattersall, 1958 G. pusillus Coifmann, 1937 G. roscoffensis Bacescu, 1970 G. sanctus (van Beneden, 1861) G. simulans W. Tattersall, 1951 G. spinifer (Goes, 1864) G. vulgaris Nakazawa, 1910 Genus Haplostylus Bacescu, 1973 H . bacescui Hatzakis, 1977 H . estafricana Bacescu, 1973 H . parerythraeus (Nouvel, 1944) Genus Iiella Bacescu, 1968 I . kojimaensis (Nakazawa, 1910) Genus Paranchialina Hansen, 1910 P. angusta (G. 0. Sam, 1883) Genus Pseudanchialina Hansen, 1910 P. erythraea Nouvel, 1944 P. inermis (Illig, 1906) P. pusilla (G. 0. Sars, 1883) P. sibogae Nouvel, 1944 Sub-family MYSINAE Tribe Erythropini Genus Amathimysis Brattegard, 1969 A. cherados Brattegard, 1974 A. gibba Brattegard, 1969 A. polita Brattegard, 1974 Genus Amblyops G. 0. Sam, 1869 A. abbreviatu (G. 0. Sars, 1869) A. aequispina Birstein and Tchindonova, 1958 A. antarctica 0. Tattersall, 1955 A. durbuni 0. Tattersall, 1955

APPENDIX I

A. ewingi Bacescu, 1967 A. kempi (Holt And Tattersall, 1905) A. magna Birstein and Tchindonova, 1958 A. spinifera Nouvel and Lagardkre, 1976 A. tattersalli Zimmer, 1914 A. tenuicazcAZa W. Tattersall, 1911 A. trisetosa Nouvel and LagardGre, 1976 Genus Amblyopsoides 0. Tattersall, 1955 A. crozetii (W.-Suhm) G. 0. Sars, 1883 A. obtusa 0. Tattersall, 1955 A. ohlinii (W. Tattersall, 1951) Genus Arachnomysis Chun, 1887 A. leuckartii Chun, 1887 A. megalops Zimmer, 1914 Genus Atlanterythrops Nouvel and Lagardbre, 1976 A. crassipes Nouvel and Lagmdhre, 1976 Genus Australerythrops W. Tattersall, 1928 A. parudicei W. Tattersall, 1928 Genus Caesaromysis Ortmann, 1893 C. hispida Ortmann, 1893 Genus Chunomysis Holt and Tattersall, 1905 C. diudema Holt and Tattersall, 1905 Genus Ductylamblyops Holt and Tattersall, 1906 D. ferwida Hansen, 1910 D. goniops W. Tattersall, 1907 D. hodgsoni Holt and Tattersall, 1906 D. iii Nouvel and Lagardhre, 1976 D. luticuuda Birstein and Tchindonova, 1958 D. latisquamsa Illig, 1960 D. murrayi W. Tattersall, 1939 D. pellucida Birstein and Tchindonova, 1958 D. sarsi (Ohlin, 1901) D. solivaga Birstein and Tchindonova, 1958 D. stenurus Murano, 1969 D. tenella Birstein and Tchindonova, 1958 D. thuumatops W. Tattersall, 1907 Genus Ductylerythrops Holt and Tattersall, 1905 D. bidigitata W. Tattersall, 1907 D. chrotqs Murano, 1969 D. dactylops Holt and Tattersall, 1905 D. dimorpha Nouvel and Lagardhre, 1976 D. gracilura W. Tattersall, 1907

327

328

THE BIOLOQY OF MYSIDS

Genus Echinomysides Murano, 1977 E. typica Murano, 1977 Genus Echinomysis Illig, 1905 E. chuni Illig, 1905 E. distinguenda Coifmano, 1936 Genus Eoerythrops Murano, 1969 E. amamiensis Murano, 1976 E . typicus Murano, 1969 Genus Erylthrops G. 0. Sam, 1869 E. abyssorum G. 0. Sara, 1869 E. africana 0. Tattersall, 1956 E. bidentata Nouvel, 1973 E. elegans (G. 0. Sam, 1863) E. erythrophthalma (Goes, 1864) E. frontieri Nouvel, 1974 E. glacialis G. 0. Sara, 1885 E. microps (G. 0. Sam, 1864) E. minuta Hansen, 1910 E. nana W. Tattersall, 1922 E. neapolitana Colosi, 1929 E. parva Brattegard, 1973 E. peterdohrni Bacescu and Schiecke, 1974 E . serrata (G. 0. Sam, 1863) E. yongei W. Tattersall, 1936 Genus Euchaetomera G. 0. Sam, 1883 E. glyphidophthalmica Illig, 1906 E. intermedia Nouvel, 1942 E. oculata Hansen, 1910 E . plebeja Hansen, 1912 E. richardi Nouvel, 1945 E. tenuis G. 0. Sam, 1883 E. typica G. 0. Sam, 1883 E . xurstrasseni (Illig, 1906) Genus Euchaetomeropsis W. Tattersall, 1909 E . merolepis (Illig, 1908) E. paciJica Banner, 1948 Genus Gibberythrops Illig, 1930 G. acunthura (Illig, 1906) G. brevisquumosa (Illig, 1906) Genus Gymnerythrops Hansen, 19 10 G. anomala Hansen, 1910 G. macrops Pillai, 1973

APPENDIX I

G. microps Coifmann, 1936 Genus Heteroerythrops 0. Tattersall, 1955 H . microps Murano, 1966 H . purpura 0. Tattersall, 1955 H . tanseii Murano, 1966 Genus Holmsiella Ortmann, 1908 H . afinis Ii, 1937 H . anomala Ortmann, 1908 Genus Hyperamblyops Birstein and Tchindonova, 1958 H . antarctica (Hansen, 1913) H . japonica (Ii, 1964) H . megalops (0.Tattersall, 1955) H . nana Birstein and Tchindonova, 1958 Genus Hypererythrops Holt and Tattersall, 1905 H . caribbaea W. Tattersall, 1937 H . elegantula Nouvel, 1974 H . richardi Bacescu, 1941 H . serriwenter Holt and Tattersall, 1905 H . spinifera (Hansen, 1910) H . zimmeri Ii, 1937 Genus Katerythrops Holt and Tattersall, 1905 K . oceanae Holt and Tattersall, 1905 K . resimora 0. Tattersall, 1955 K . tattersalli Illig, 1930 K . triangulata Panarnpunnayil, 1977 Genus Longithorax Illig, 1906 L. alicei Nouvel, 1942 L. capensis Zknmer, 1914 L. fuscus Hansen, 1908 L. nouveli 0. Tattersall, 1955 L. similerythrops Illig, 1906 Genus Metamblyops W. Tattersall, 1907 M . macrops W. Tattersall, 1937 M . oculata W. Tattersall, 1907 M . philippinensis (W. Tattersall, 1951) M . Stephenson; W. Tattersall, 1936 Genus Meterythrops S. I. Smith, 1879 M . japonica Murano, 1977 M . megalops Ii, 1964 M . microphthalma W. Tattersall, 1951 M . picta Holt and Tattersall, 1905 M . robusta S. I. Smith, 1879

329

330

THE BIOLOGY OF MYSIDS

Genus Michthyops W. Tattersall, 1911 M . parwa (Vanhoffen, 1897) M . theeli (Ohlin, 1901) Genus Nysimenzies Bacescu, 197 1 M . hadalis Bacescu, 1971 Genus Nipponerythrops Murano, 1977 N . typica Mkano, 1977 Genus Paramblyops Holt and Tattersall, 1905 P. bidigitata W. Tattersall, 1911 P. brevirostris 0. Tattersall, 1955 P. globorostris Birstein and Tchindonova, 1970 P. rostrata Holt and Tattersall, 1905 Genus Parapseudomma Nouvel and Lagardhe, 1976 P. calloplura (Holt and Tattersall, 1905) Genus Parerythrops G. 0. Sars, 1869 P. afinis Birstein and Tchindonova, 1958 P. bispinosa Nouvel and Lagardbre, 1976 P. lobiancoi W. Tattersall, 1909 P. obesa (G. 0. Sars, 1864) P. paucispinosa Nouvel and Lagardhre, 1976 P. spectabilis G. 0. Sars, 1885 Genus Pleurerythrops Ii, 1964 P. constricta Panampunnayil, 1977 P. inscita Ii, 1964 P. secunda Murano, 1970 Genus Pseudamblyops Ii, 1964 P . conicops Ii, 1964 Genus Pseuderythrops Coifmann, 1936 P. gracilis Coifmann, 1936 Genus Pseudomma G. 0. Sars, 1870 P. afine G. 0. Sars, 1870 P. antarcticum Zhmer, 1914 P. armatum Hansen, 1913 P. australe (G. 0. Sara, 1883) P. belgicae (Hansen in MS) Holt and Tattersall, 1906 P. berkeleyi W. Tattersall, 1933 P. bispinicaudurn Murano, 1974 P. brevisqwmosum Murano, 1974 P. calmani 0. Tattersall, 1955 P. chuttoni Bacescu, 1941 P. crassidentatum Murano, 1974 P. frigidum Hansen, 1908

APPENDIX I

P. intermedium Murano, 1974 P. izuensis Murano, 1966 P. japonicum Murano, 1970 P. kruppi W. Tattersall, 1909 P. lamellicaudum Murano, 1974 P. latiphthalmum Murano, 1974 P. longicaudum 0. Tattersall, 1955 P. longisquamosum Murano, 1974 P. magellanensis 0. Tattersall, 1955 P. marumoi Murano, 1974 P. matsuei Murano, 1966 P. minutum 0. Tattersall, 1955 P. multispina Birstein and Tchindonova, 1958 P. nanum Holt and Tattersall, 1906 P. okiyamai Murano, 1974 * P. omoi Holmquist, 1957 P. roseum G. 0. Sam, 1870 P. sarsii (W.-Suhm in MS) G. 0. Sam, 1883 P. schollaertensis 0. Tattersall, 1955 P. surugae Murano, 1974 P. tameii Murano, 1974 P. truncatum S. I. Smith, 1879 Genus Pteromysis Ii, 1964 P. amemiyai Ii, 1964 Genus Scolamblyops Murano, 1974 S. japonicw Murano, 1974 S. oculospinum (W. Tattersall, 1951) Genus Synerythrops Hansen, 1910 8.cruciata W. Tattersall, 1951 8.intermedia Hansen, 1910 S. truncuta Murano, 1975 Genus Teraterythrops Ii, 1964 T . parva (Zimmer, 1914) T . robusta (Birstein and Tchindonova, 1958) Genus Thlassomysis W. Tattersall, 1939 T . sewelli W. Tattersall, 1939 T . tattersalli Nouvel, 1942 Tribe Leptomysini Genus Afromysis Zimmer, 1916 A. bainbridgei 0. Tattersall, 1957 A . dentisinus Pillai, 1957 A . guinensis Bacescu, 1968

331

332

THE BIOLOGY OF MYSIDS

A . hansoni Zimmer, 1916 A . macropsis W. Tattersall, 1922 A . ornata 0. Tattersall, 1957 Genus Australomysis W. Tattersall, 1927 A . acuta W. Tattersall, 1927 A . incisa (G. 0. Sars, 1883) Genus Bathymysis W. TaDtersall, 1907 B. helgae W. Tattersall, 1907 B. renoculata W. Tattersall, 1951 Genus Brasilomysis Bacescu, 1968 B. castroi Bacescu, 1968 B. inermis (Coifmann, 1937) Genus Calyptomma W. Tattersall, 1909 G. puritani W. Tattersall, 1909 Genus Cubanomysis Bacescu, 1968 C. jimenesi Bacescu, 1968 Genus Dioptromysis Zimmer, 1915 D. paucispinosa Brattegrad, 1969 D. perspicillata Zimmer, 1915 D. proxima Nouvel, 1964 D . spinosa Brattegard, 1969 Genus Doxomysis Hansen, 1912 D. anomala W. Tattersall, 1922 D . australiensis (W. Tattersall, 1940) D. hanseni Colosi, 1920 D . littoralis W. Tattersall, 1922 D . longiura Pillai, 1963 D. microps Colosi, 1920 D. quadrispinosa (Illig, 1906) D. zimmeri Colosi, 1920 Genus Hyperiimysis Nouvel, 1966 H . madagascariensis Nouvel, 1966 Genus Iimysis Nouvel, 1966 I . atlantica (Nouvel, 1942) 1. orientalis (Ii, 1937) Genus Leptomysis G. 0. Sars, 1869 L. apiops G. 0. Sam, 1877 L. australiensis W. Tattersall, 1927 L. burgii Bacescu, 1966 L. capensis Illig, 1906 L. gracilis (G. 0. Sars, 1864) L. lingvura (G. 0. Sars, 1866)

APPENDIX I

L. mediterranea G. 0. Sars, 1877 L. megalops Zhmer, 1915 L. peresi Bacescu, 1966 L. sardica G. 0. Sars, 1877 L. xenops W. Tattersall, 1922 Genus Metamysidopsis W. Tattersall, 1951 M . elongata (Holmes, 1900) M . insularis Brattegard, 1970 M . macaensis Silva, 1970 M . mexicana Bacescu, 1969 M . munda (Zimmer, 1918) M . pacifica (Zimmer, 1918) M . swifti Bacescu, 1969 Genus Mysideis G. 0. Sars, 1869 M . insignis (G. 0. Sars, 1864) M . parva Zimmer, 1915 Genus Mysidetes Holt and Tattersall, 1906 M . anomala 0. Tattersall, 1955 M . antarctica 0. Tattersall, 1965 M . brachylepis W. Tattersall, 1923 M . crassa Hansen, 1913 M . dimorpha 0. Tattersall, 1955 M . farrani (Holt and Tattersall, 1905) M . hanseni Zimmer, 1914 M . intermedia 0. Tattersall, 1955 M . kerguelensis (Illig, 1906) M . macrops 0. Tattersall, 1955 M . microps 0. Tattersall, 1955 M . patagonica 0. Tattersall, 1955 M . peruana Bacescu, 1967 M . posthon Holt and Tattersall, 1906 Genus Mysidopsis G. 0. Sars, 1864 M . acuta Hansen, 1913 M . almyra Bowman, 1964 M . angusta G. 0. Sars, 1864 M . ankeli Brattegard, 1973 *. M . arenosa Brattegard, 1974 M . bahia Molenock, 1969 M . bigelowi W. Tattersall, 1927 M . bispinosa 0. Tattersall, 1969 M . bispinulata Brattegard, 1974 M . brattstroemi Brattegard, 1969

333

334

THE BIOLOGY OF MYSIDS

M . californica W. Tattersall, 1932 M . camelin& 0. Tattersall, 1955 M . coelhoi Bacescu, 1968 M . coralicola Bacescu, 1975 M . cultrata Brattegard, 1973 M . didelphys (Norman, 1863) M . eclipes Brattegard, 1969 M . eremita 0. Tattersall, 1962 M . furca Bowman, 1957 M . gibbosa G. 0. Sars, 1864 M . hellvillensis Nouvel, 1964 M . indica W. Tattersall, 1922 M . intii Holmquist, 1957 M . japonica Ii, 1964 M . kempi W. Tattersall, 1922 M . kenyana Bacescu and Vasilescu, 1973 M . major (Zirnmer, 1912) M . mathewsoni Brattegard, 1969 M . mauchlinei Brattegard, 1974 M . mortenseni W. Tattersall, 1951 M . robusta Brattegard, 1974 M . robustispina Brattegard, 1969 M . schultzei (Zimmer, 1912) M . similis (Zimmer, 1912) M . suedafrikana 0. Tattersall, 1969 M . surugae Murano, 1970 ..W.taironana Brgttegard, 1973 M . tortonesei Bacescu, 1968 M . velifera Brattegard, 1973 M . virgulata Brattegard, 1974 Genus Nouvelia Bacescu and Vasilescu, 197 3 N . natalensis (0.Tattersall, 1952) N . nigeriensis (0.Tattersall, 1957) N . valdiviae (Illig, 1906) Genus Prionomysk W. Tattersall, 1922 P. aspera Ii, 1937 P. stenolepis W. Tattersall, 1922 Genus Promysis Dana, 1850 P. atlantica W. Tattersall, 1923 P. orientalis Dana, 1852 Genus Pseudomysidetes W. Tattersall, 1936 P. cochinensis Panampunnayil, 1977

APPENDIX I

P . russelli W. Tattersall, 1936 Genus Pseudomysis G. 0. Sars, 1879 P. abyssi G. 0. Sars, 1885 P . dactylops W. Tattersall, 1951 Genus Pseudoxomysis Nouvel, 1973 P. caudaensis Nouvel, 1973 Genus Tenagomysis Thomson, 1900 T . chiltoni W. Tattersall, 1923 T . macropsis W. Tattersall, 1923 T . novae-zealandiae Thomson, 1900 T . producta W. Tattersall, 1923 T . robusta W. Tattersall, 1923 T . scotti W. Tattersall, 1923 T . similis W. Tattersall, 1923 T . tanzaniana Bacescu, 1975 T . tenuipes W. Tattersall, 1918 T . thomsoni W. Tattersall. 1923 Tribe Mysini Genus Acanthomysis Czerniavsky, 1882 A. alaskensis Banner, 1954 A. anomala Pillai, 1961 A . aokii Ii, 1964 A. aspera Ii, 1964 A. borealis Banner, 1954 A . columbiae (W. Tattersall, 1933) A . costata (Holmes, 1900) A. davisi Banner, 1948 A. dimorpha Ii, 1936 A . dybowskii (Derjavin, 1913) A. fujinagai Ii, 1964 A . grimmi (G. 0. Sam, 1895) A . hodgarti (W. Tattersall, 1922) A . hwanhaiensis Ii, 1964 A. Bndica (W. Tattersall, 1922) A . inJlata (G. 0. Sam, 1907) A. japonjca (Marukawa, 1928) A. koreana Ii, 1964 A . longicornis (M.-Edwards, 1837) A. longirostris Ii, 1936 A. macrops Pillai, 1973 A . macropsis (W. Tattersall, 1932) A . mitsukurii (Nakazawa, 1910)

335

336

THE BIOLOGY OF MYSIDS

A. nakazatoi ,Ii, 1964 A. nephrophthalma Banner, 1948 A. okayamaensis Pi, 1964 A . ornata 0. Tattersall, 1965 A. pelagica (Pillai, 1957) A . platycauda (Pillai, 1964) A . pseudomacropsis (W. Tattersall, 1933) A. pseudomitsukurii Ii, 1964 A. quadrispinosa Nouvel, 1965 A. sagamiensis (Nakazawa, 1910) A. schrencki (Czerniavsky, 1882) A. sculpta (W. Tattersall, 1933) A. sinensis Iit1964 A. stelleri (Derjavin, 1913) A. strauchi (Czerniavsky, 1882) A. tamurai Ii, 1964 A . trophopristes 0. Tattersall, 1957 Genus Anisomysis Hansen, 1910 A. aikuwai Ii, 1964 A. bifhrcata W. Tattersall, 1912 A. bipartoculata Ii, 1964 A . hanseni Nouvel, 1967 A. hispida Pillai, 1973 A. ijimlai Nakazawa, 1910 A. incisa W. Tattersall, 1936 A. kunduchiana Bacescu, 1975 A. lamellicauda (Hansen, 1912) A. laticauda Hansen, 1910 A. levi Bacescu, 1973 A. maris rubri Bacescu, 1973 A. megalops (Illig, 1913) A. mixta Nakazawa, 1910 A. ielewensis Ii, 1964 A. sirielloides Bacescu, 1975 A. tattersallae Pillai, 1973 A. thurneysseni Nouvel, 1973 A. vmseuri Ledoyer, 1974 Genus Antarctomysis Coutihre, 1906 A. maxima (Hansen) Holt and TatCeniall, 1906 A. ohlinii Hansen, 1908 Genus Antromysis Creaser, 1936 A. almyra (Brattegard, 1977)

APPENDIX I

A . americana (W. Tattersall, 1951) A . anophelinae W. Tattersall, 1951 A . bahumensis (Brattegard: 1969) A . cenotensis Creaser, 1936 A . cubanica Bacescu and Orghidan, 1971 A . juberthiei Bacescu and Orghidan, 1977 A . peckorum Bowman, 1977 A . reddelli Bowman, 1977 Genus Arthromysis Colosi, 1924 A . magellanica (Cunningham, 1871) Genus Canegieomysis W. Tattersall, 1943 C . xenqs W. Tattersall, 1943 Genus Caspiomysis G. 0. Sars, 1907 C. knipowitschi G. 0 . Sara, 1907 Genus Diamysis Czerniavsky, 1882 D. bahirensis (G. 0 . Sam, 1877). D . frontieri.Nouve1, 1965 D. pengoi (Czerniavsky, 1882) D.pusilla (G. 0. Sam, 1907) Genus Gangemysis Tattersall and Tattersall, 1951 G. assirnilis (W. Tattersall, 1908) Genus Hemimysis G. 0. Sars, 1869 H . abyssicola G. 0 . Sars, 1869 H . anomala G. 0 . Sars, 1907 H . lamornae (Couch, 1856) H . serrata Bacescu, 1938 H . speluncola Ledoyer, 1963 Genus Idiomysis W. Tattersall, 1922 I . inermis W. Tattersall, 1922 I . japonica Murano, 1978 I . kurnamali Bacescu, 1973 Genus Indomysis W. Tattersall, 1914 I . annanhlei W. Tattersall, 1914 Genus Inusitatomysis Ii, 1940 I . insolita Ii, 1940 Genus Kainomatomysis W. Tattersall, 1927 K. foxi W. Tattersall, 1927 K . schieckei Bacescu, 1973 Genus Katamysis G. 0. Sars, 1877 K. warpachowskyi G. 0. Sars, 1877 Genus Limnomysis Czerniavsky, 1882 L.benedeni Czernaivsky, 1882

337

338

THE BIOLOGY OF MYSIDS

Genus Lycomysis Hansen, 1910 L. bispina Ii, 1940 L. platycauda Pillai, 1961 L. spinicauda Hansen, 1910 Genus Mesacanthomysis Nouvel, 1967 M . pygmaea Nouvel, 1967 Genus Mesopodopsis Czerniavsky, 1882 M . africana 0. Tattersall, 1952 M . orientalis (W. Tattersall, 1908) M . slabberi (van Beneden, 1861) M . zeylanica Nouvel, 1954 Genus Mysidium Dana, 1850 M . columbiae (Zimmer, 1915) M . gracile (Dana, 1852) M . integrum W. Tattersall, 1951 Genus Mysis Latreille, 1803 M . amblyops G. 0. Sars, 1907 M . aqstrale 0. Tattersall, 1955 M . caspia G. 0. Sam, 1895 M . gaspensis 0. Tattersall, 1954 M . litoralis (Banner, 1948) M . macrolepis G. 0. Sars, 1907 M . microphthulma G. 0. Sam, 1895 M . mista Lilljeborg, 1852 M . oculata (Fabricius, 1780) M . polaris Holmquist, 1959 M . relicta LovBn, 1862 M . stenolepis S. I. Smith, 1873 Genus Nanomysis W. Tattersall, 1921 N . insularis Nouvel, 1957 N . siamensis W. Tattersall, 1921 Genus Neomysis Czerniavsky, 1882 N . americana (S. 1. Smith, 1874) N . awatschensis (Brandt, 1851) N . czerniavskii Dershavin, 1913 N . ilyapai Holmquist, 1957 N . integer (Leach, 1814) N . intermedia (Czerniavsky, 1882) N . japonica Nakazawa, 1910 N . kadiakensis Ortmann, 1908 N . mercedis Holmes, 1896 N . meridionalis Colosi, 1924

AF'PENDIX I

N . mirabilis (Czerniavsky, 1882) N . monticeltii Colosi, 1924 N . orientalis Ii, 1964 N . patagona Zimmer, 1907 N . rayii (Murdoch, 1885) N . sopayi Holmquist, 1957 N . spinosa Nakazawa, 1910 Genus Paracanthomysis Ii, 1936 P. hispida Ii, 1936 P. kurilensis Ii, 1936 Genus Paramysis Czerniavsky, 1882 P. agigensis Bacescu, 1938 P. arenosa (G. 0. Sars, 1877) P. bacescoi Labat, 1953 P. baeri Czerniavsky, 1882 P. bakuensis G. 0. Sam, 1895 P. eurylepis G. 0. Sars, 1907 P. festae Colosi, 1920 P. grimmi (G. 0. Sam, 1895) P. helleri (G. 0. Sam, 1877) P. incerta (G. 0. Sars, 1895) P. inflata (G. 0. Sam, 1907) P. intermedia (Czerniavsky, 1882) P. kessleri (Grimm, 1875) P. kosswigi Bacescu, 1948 P. krnyeri (Czerniavsky, 1882) P. lacustris (Czerniavsky, 1882) P. Zoxolepis (G. 0. Sars, 1895) P. nouveli Labat, 1953 P. pontica (Bacescu, 1938) P. portxicensis Nouvel, 1950 P. proconnesia Colosi, 1922 P. ullskyi (Czerniavsky, 1882) Genus Parastilomysis Ii, 1936 P. paradoxa Ii, 1936 Genus Praunus Leach, 1814 P. flexuosus (Miiller, 1776) P. inermis (Rathke, 1843) P. neglectus (G. 0. Sam, 1869) Genus Proneomysis W. Tattersall, 1933 P. eriopedeu Ii, 1936 P. fusca Ii, 1936

339

340

THE BIOLOGY OF MYSIDS

P. lingvura Murano, 1977 P. longipes Murano, 1977 P. misakiensis Ii, 1936 P.ornata Ii, 1964 P. perminuta Ii, 1936 P.quadrispinosa Ii, 1964 P.sandoi Ii, 1963 P. surugensis Murano, 1977 P. takitai Murano, 1977 P. tenuiculus Ii, 1940 P. toriumii Murano, 1977 P. wailesi W. Tattersall, 1933 Genus Schistomysis Norman, 1892 8. assimilis (G. 0. Sars, 1977) S. elegans G. 0. Sars, 1907 S. kervillei (G. 0. Sars, 1885) 8. ornata (G. 0. Sam, 1864) S. parkeri Norman, 1892 8. spiritus (Norman, 1860) Genus Stilomysis Norman, 1892 S. camtschatica Marukawa, 1928 X. grandis (Goes, 1863) 8. major W. Tattersall, 1951 Genus Taphromysis Banner, 1953 T . bowmani Bacescu, 1961 T.louisianae Banner, 1953 Genus Troglomysis Stammer, 1933 T . vjetrenicensis Stammer, 1933 Tribe Heteromysini Genus Heteromysis S. I. Smith, 1874 H . actiniae Clarke, 1955 H. armoricana Nouvel, 1940 H . atlantidea 0. Tattersall, 1961 H. bermudensis G. 0. Sam, 1885 H . bredini Brattegard, 1970 H. brucei 0. Tattersall, 1967 H . digitata W. Tattersall, 1927 H . dispar Brattegard, 1970 H . disrupta Brattegard, 1970 H . eideri Bacescu, 1941 H. elegans Brattegard, 1974 H . $oridensis Brattegard, 1969

APPENDIX I

H:formosa S. I. Smith, 1874 H . gerlmhei (Bohnier and Perez, 1902) H . qomezi Bacescu, 1970 H . quitarti Bacescu, 1968 H . gymnura W. Tattersall, 1922 H . harpax (Hilgendorf, 1879) H. kossmanni Nouvel, 1964 H . lybiana Bacescu, 1976 H . macropsis Pillai, 1961 H . mariani Bacescu, 1970 H . mayana Brattegard, 1970 H . microps (G. 0. Sam, 1877) H . minuta 0. Tattersall, 1967 H . nouveli Brattegard, 1969 H . ohntops Walker, 1898 H . pmi$ca 0. Tattersall, 1967 H . panamaensis 0. Tattersall, 1967 H . proxima W. Tattersall, 1922 H. rubrocincta Bacescu, 1968 H . siciliseta Brattegard, 1970 H . sinqaporensis 0. Tattersall, 1967 H . tasmanica W. Tattersall, 1927 H . tattersalli Nouvel, 1942 H . waitei W. Tattersall, 1927 H . xanthops Ii, 1964 H . zeylanica W. Tattersall, 1922 Genus Heteromysoides Bacescu, 1968 H . cotti (Calman, 1932) H . spongicola Bacescu, 1968 Genus Neoheteromysis Bacescu, 1976 N . mfilleri Bacescu, 1976 Sub-family MYSIDELLINAE Genus Mysidella G. 0. Sam, 1872 M . americana Banner, 1948 M . minuta Brattegard, 1973 M . nana Murano, 1970 M . tanakai Ii, 1964 M . typhlops G. 0. Sars, 1872 M . typica G. 0. Sam, 1872 Family LEPIDOMYSIDAE Genus Spelaeomysiv Caroli, 1924 8.bottazzii Caroli, 1924

341

342

THE BIOLOGY OF MYSIDS

S. cardisomae Bowman, 1973 S . longipes (Fillai and Mariamma, 1964) S. nuniezi Bacescu and Orghidan, 1971 S. olivae Bowman, 1973 S. quinterensis (Villalobos, 1951) S. servatus (Fage, 1924) Family STYGIOMYSIDAE Genus Stygiomysis Caroli, 1937 S. holthuisi (Gordon, 1960) S. hydruntina Caroli, 1937 S. major Bowman, 1976

APPENDIX I1

CLASSIFIED LIST OF LITERATURE FOR EACH GENUS Gordan (1957), in her bibliography of the Mysidacea, classifies the literature according to species but this method is too cumbersome for this appendix. She closed her bibliography in 1955 and the subsequent literature, as well as some references omitted by her, are cited here for each genus. The genera are listed alphabetically, as they are in the World list of species published by Mauchline and Murano (1977). The broad geographical occurrence of each species was listed in the World List but no sources of information were given. These sources are now published in this appendix.

Amnthomysis Ostenfeld, 1906; Ostenfeld and Wesenberg-Lund, 1909 ; Herdman, 1920; Bacescu, 1936a, 1975; Banner, 1954a; Raw, 1955; Anonymous, 1967; Nouvel, 1951a, 1965; Caspers, 1957; 0. S. Tattersall, 1957, 1958, 1960a, b, 1965a; Bainbridge, 1960; Pillai, 1961, 1964, 1965, 1973; Wickstead, 1961; Hoenigman, 1963, 1968; Balasubrahmanyan, 1964; Ii, 1964s; Wing, 1965,1975; Hoffman and Yancey, 1966; Painter, 1966; Ariani, 1967; Clutter, 1967; Haertel and Osterberg, 1967; Kikuchi, 1967; Taniguchi, 1969; Champelbert and Macquart-Moulin, 1970; Elofsson and Dahl, 1970; Green, 1970; Field, 1971; Friedl, 1971a, b; Mauchline, 19718, f, g, 197313; Ghirardelli, 1973; Vader, 1973; Simmons et al., 1974; Hobson and Chess, 1976; Makings, 1977; Mauchline and Murano, 1977; Wing and Barr, 1977; Wittmann, 1977; Omori, 1978.

Afromysis Colosi, 1923; Dakin and Colefax, 1940; 0. S. Tattersall, 1957, 1958, 1961b, 1965a; Bainbridge, 1960; Balasubrahmanyan, 1964; Pillai, 1964, 1965, 1973; Nouvel, 1966s; Bacescu, 19688; Mauchline, 197313; Shyamasundari, 1973; Mauchline and Murano, 1977.

Amathimpis Brattegard, 1969, 1970b, 1973, 19748, b, 1975; Mauchline and Murano, 1977.

344

THE BIOLOGY OF MYSIDS

A mblyaps Ostenfeld and Wesenberg-Lund, 1909; Banner, 19548,; Birstein and Tchindonova, 1958, 1970; Fisher and Kon, 1959; Ii, 1964a; 0. S. Tattersall, 1965; Belyaev, 1966; Bacescu, 1967; O’Riordan, 1969; Wigley and Burns, 1971; Mauchline, 1973b; Nouvel and Lagardhe, 1976; Elofsson and Hallberg, 1977; Lagarddre, 1977b; Mauchline and Murano, 1977.

Amblyopoides Holmquist, 1957b; 0. S. Tattersall, 1965b; Mauchline and Murano, 1977.

Anchialina Scott, 1900a ; Ostenfeld, 1906, 1916 ; Ostenf‘eld and Wesenberg-Lund, 1909; Zimmer, 1912; Dakin and Colefax, 1940; Nouvel, 1944a, 1951a, 1969, 1971; Fraser and Saville, 1949a, b; Banner, 1954a; Hoenigman, 1954, 1960, 1963, 1968; Le Sueur, 1954; Anonymous, 1957; Furnestin, 1959; Holmquist, 1959b, Reys, 1960; 0. S. Tattersall, 1960a, 196lb, 1962, 1965a; Evans, 1961; F’raser, 1961, 1970; Wickstead, 1961; Haidar, 1963; Ii, 1964a; Pillai, 1964, 1965, 1973; Macquart-Moulin, 1965, 1972, 1973b, 1977b; Hopkins, 1966; Ledoyer, 1966; Ariani, 1967 Vives, 1968; O’Riordan, 1969; Brattegard, 1970a, 1973,1975; Champelbert and Macquart-Moulin, 1970; Baan and Holthuis, 1971;Mauchline, 1971e, f, g, 1972, 1973b, c; Wigley and Burns, 1971; Lagardhre, 1972 1977a, b ; Ghirardelli, 1973; Vader, 1973; Haezakis, 1974; Koirtsis, 1974; Bacescu, 1975, 1976; Ariani and Spagnwla, 1976; Bacescu and Muradian, 1977; Beaudouin, 1977; Makings, 1977; Mauchline and Murano, 1977; Silva, 1977; Wittmann, 1977.

Anisomysis Coutihre, 1917; Ii, 1964a; Nouvel, 1967, 1973b; Bacescu, 1973b, c, d, 1975; Mauchline, 197313; Pillai, 1973; Ledoyer, 1974; Mauchline and Murano, 1977; Pratap et al., 1977.

Antarctomysis Rustad, 1930; Mackintosh, 1934; Boschma, 1949; Holmquist, 1959b; 0. S. Tattersall, 1961a, 1965b; Birstein and Tchindonova, 1962; Ledoyer, 1969; Mauchline, 1972, 1973b; Mauchline and Murano, 1977.

Antromysis Brattegard, 1969, 1970b, 1973, 1974a, b, 1975, 1977 (Parvimysis); Bacescu and Orghidan, 1971; Mauchline, 1973b; Bowman, 1977a, b; Mauchline and Murano, 1977; Crouau, 1978.

APPENDIX II

345

A rachnomysis C m t i , 1921; Fraser, 1961; Hoenigman, 1963; Pillai, 1973; Mauchline and Murano, 1977; Murano, 1977a.

A rchueomysis Banner, 1954a; Enright, 1962; Ii, 1964a; Elofsson, 1965; Clutter, 1967; Haertel and Osterberg, 1967; Johnson, 1969; Jawed, 1973; Kasaoka, 1974; Holmquist, 1975; Mauchline and Murano, 1977.

A rthromysis Holmquist, 1957b; 0 . S. Tattersall, 1965b; Mauchline and Murano, 1977.

Atlvnterythrops Nouvel and Lagardhre, 1976; Lagardhre, 197713; Mauchline and Murano, 1977.

AuatraZerythrops Mauchline and Murano, 1977. A uatralomysis Dakin and Colefax, 1940; Mauchline and Murano, 1977: Bell et al., 1978.

Bathymysis Pillai, 1963b, 1964, 1965; O’Riordan, 1969; Murano, 1970c; Wigley and Burns, 1971; Lagardhre, 1977b; Mauchline and Murano, 1977.

Boreomysis Norman, 1880b; Hjorti and Dahl, 1900; Ostenfeld, 1906; Ostenfeld and Wesenberg-Lund, 1909; Pawsey and Davis, 1924; Stephensen, 1933,1943; Hanstrom, 1948; Banner, 1954a; Ostvedt, 1955; Holmquist, 1956, 1957a, b ; Birstein and Tchindonova, 1958, 1962; Belloc, 1959; Tchindonova, 1959; Brunel, 1960, 1961, 1965, 1970; Gordon, 1960; Bacescu and Mayer, 1961; Clarke, 1961a; Fraser, 1961; 0. S. Tattersall, 1961b, 1965b; F’refontaine and B m e l , 1962; Vinogradov, 1962; Drainville et al., 1963; Hoenigman, 1963; Ii, 1964a; Pillai, 1964, 1965, 1973; Elofsson, 1965; Jepsen, 1965; Dahl and Mecklenburg, 1969; Geiger, 1969; O’Riordan, 1969; Taniguchi, 1969; Casanova, 1970; Zharkova, 1970; Childress, 1971b, 1975; Wiborg, 1971; Wigley and Burns, 1971; Lagardhre, 1972, 1977a, b ; Mauchline, 1972, 197313; Nemoto, 1972; Childress and Nygaard, 1974; Haedrich and Henderson, 1974; Judkins and Wright, 1974; Murano et al., 1976; Elofsson and

346

THE BIOLOGY OF MYSIDS

Hallberg, 1977; Matthews and Bakke, 1977; Mauchline and Murano, 1977; Pearcy et al., 1977; Slijoldal and Bamstedt, 1977; Bamstedt, 1978; Shih and Laubitz, 1978.

Bowmaniella Bacescu, 1968d, 1969; Brattegard, 1970a, b, 1973, 1974a, b, 1975; Sib, 1970, 1971a, b, 1972; Wigley and Burns, 1971; Almeida Prado, 1972, 1973, 1974; Williams, 1972; Mauchline, 1973b; Holmqujst, 1975; Holland and Polgar, 1976; Camp et al., 1977; Mauchline and Murano, 1977; Silva, 1977; Price, 1978. Brasilomysis Bacescu, 1968a, 1969; Brattegard, 1969, 1973, 1974a, b ; Almeida Prado, 1972, 1973, 1974; Mauchline, 1973b; Mauchline and Murano, 1977; Silva, 1977. Caesaromysis Stebbing, 1905; Zimmer, 1912; Banner, 1954a; Birstein and Tchindonova, 1958,1962; Mauchline and Murano, 1977; Murano, 1977a. Calyptommu MauchIine and Murano, 1977 Carnegieomysis Mauchline and Murano, 1977. Caspiornysis Mauchline and Murano, 1977. Ceratolepis Coutihre, 1904; Clarke, 1961a; Mauchline, 1973b; Mauchline and Mwano, 1977. Ceratornysis Birstein and Tchindonova, 1958; Ledoyer, 1977; Mauchline and Murano, 1977. Chalaraspidum Banner, 1954a; Clarke, 1961a; Birstein and Tchindonova, 1962; 0. S. Tattersall, 196513; Holthuis and Sivertsen, 1967; Mauchline, 1972; Mauchline and Murano, 1977.

APPENDIX II

347

Chummysis Osbnfeld and Wesenberg-Lund, 1909; O’Riordan, 1969; Nouvel and LagardBre, 1976; Mauchline and Murano, 1977.

Cubanomysis Bacescu, 1968c; Brattegard, 1973, 19748, b, 1975; Mauchline, 1973b; Mauchline and Murano, 1977.

Dactylamblyops Birstein and Tchindonova, 1958, 1962; Tchindonova, 1959; Ii, 1964a; Pillai, 1964, 1965; 0.S. Tattersall, 1965b; Murtmo, 1969; O’Riordan, 1969; Zharkova, 1970; Nemoto, 1972; Nouvel and LagardBre, 1976; Lagardhre, 1977b; Mauchline and Murano, 1977.

Dactylerythrqs Ostenfeld and Wesenberg-Lund, 1909;Murano, 1969,1970~; O’Riordan, 1969; Nouvel and LagardBre, 1976; Lagmdhre, 1977b; Mauchline and Murmo, 1977.

Diamysis Bacescu, 1936a, 1938, 1954, 1973d, 1975; Gemvese, 1956; Valkanov, 1957;Ben’ko, 1962; Holmquist, 1962; Hoenigman, 1963, 1968; Nouvel, 1965; Zatkutskiy, 1965,1970; Dediu, 1966;Mauchline, 1973b; Bowman, 1977b ~ ;~Brattegard, 1977; Mauchline and Murano, 1977.

Dioptromysis Pillai, 1963d, 1964, 1965; Nouvel, 1964a; Brattegard, 1969; Ledoyer, 1970; Bacescu, 1975; Mauchline and Murano, 1977.

Doxomysis Colosi, 1923; Pillai, 1963a, 1964, 1965, 1973; Ii, 1964a; Nouvel, 1966a; Taniguchi, 1976; Mauchline and Murano, 1977. Echimmysioks Mauchline and Murtmo, 1977 ; Murano, 1977a. Echinomysis F’illai, 1964, 1965; Mauchline and Murmo, 1977.

Eoerythrops Murrtno, 1969, 1970c, 1976; Mauchline and Murano, 1977.

348

THE BIOLOGY OF MYSIDS

Erythrops T . Scott, 1887, 1889, 1890, 1896b, 1897, 1898, 1900a, 1902a, b ; Walker, 1898b; Ostenfeld, 1906, 1916; Ostenfeld and Wesenberg-Lund, 1909; Herdman, 1920; Moore, 1937; Stephensen, 1943; Whiteley, 1948; Jones, 1954; Anonymous, 1957; Beyer, 1958; Fisher and Kon, 1959; Furnestin, 1959; Brunel, 1960, 1961, 1963a, 1970; Reys, 1960; 0. S . Tattersall, 1960a, 1965; Clarke, 1961a; Fraser, 1961; Wickstead, 1961; Drainville et al., 1963; Balasubrahmanyan, 1964; Ii, 1964a; Pillai, 1964, 1965, 1973; Macquart-Moulin, 1965; Nagabhushanam, 1965; Hoenigman, 1968; Mauchline, 1968, 1969b, 1971f, g, 1972, 1973a, b, c, 1977a; O’Riordan, 1969; Poirier,l969; Champelbert and MacquartMoulin, 1970; Elofsson and Dahl, 1970; Just, 1970; Wigleyand Burns, 1971; Brattegard and Vader, 1972; Brattegard, 1973,1974a; Hesthagen, 1973; Nouvel, 1973d, 1974; Shyamasundari, 1973; Bacescu and Schiecke, 1974; McEachran et al., 1976; Bacescu and Muradian, 1977; Beaudouin, 1977; Hallberg, 1977; Lagardkre, 1977b; Makings, 1977; Mauchline and Murano, 1977; Hesthagen and Gjermundsen, 1978; Hopkins and Gulliksen, 1978; Schnack, 1978.

Euchuetomera Colosi, 1923; Rustad, 1930; Banner, 1954a; Belloc, 1959; Birstein and Tchindonova, 1962; 0. S. Tattersall, 1962, 1965b; Ii, 1964s; Pillai, 1964, 1965, 1973; Nouvel and Lagardere, 1976; Taniguchi, 1976; Mauchline and Murano, 1977; Murano, 19778.

Euchaetomeropsis Banner, 1954a; Casanova, 1968, 1970; Taniguchi, 1976; Mauchline and Murano, 1977; Murano, 1977a.

Eucopia Richard, 1904; Stebbing, 1914; Nouvel, 194413; Hanstrom, 1948; Fraser, 1950, 1956, 1961; Siewing, 1953; Banner, 1954a; Raw, 1955; Bruun, 1957; Birstein and Tchindonova, 1958, 1962; Belloc, 1959; Tchindonova, 1959; Brunel, 1960; Dion and Nouvel, 1960; Furnestin, 1960; Gordon, 1960; Clarke, 1961a; 0. S. Tattersall, 1961b, 1965b; Hoenigman, 1963; Elofsson, 1965; Macquart-Moulin, 1965; MacquartMoulin and Leveau, 1968; Vives, 1968; O’Riordan, 1969; Raymont et al., 1969; Taniguchi, 1969, 1976; Casanova,, 1970, 1977; Elofsson and Dahl, 1970; Vinogradov, 1970; Zharkova, 1970; Franqueville, 1971; Wigley and Burns, 1971; Mauchline, 1972, 1973b, 1977a; Morris, 1972;

APPENDIX 11

349

Herring, 1974; Murano, et al., 1976; Lagardbre, 1977a, b; Mauchline and Murano,. 1977; Moore and Sander, 1977; Pearcy et al., 1977; Sargent et at., 1978.

Qangemysis Gordan (1957) as Potamomysis arntrosmcus Walker, 1888b, 1898b; Herdman, 1890; Herdman et al., 1898; Scott, 1902b; Ostenfeld, 1906, 1916; Gilson, 1909; Ostenfeld and WesenbergLund, 1909; Petersen, 1914; Todd, 1914; Hardy, 1924; Savage, 1931; Moore, 1937; Bacescu, 1938, 1954, 1968d, 1970a, 1975, 1976; Rees, 1939; Dakin and Colefax, 1940; Nouvel, 1944a, 1951a, b, 1973c; Matsudaira et al., 1952; Delamare Deboutteville, 1953a, b, c; Jones, 1954; Le Sueur, 1954; Taberly, 1954a, b; Colman and Segrove, 1955; Anonymous, 1957; Suau and Vives, 1957; 0. S. Tattersall, 1957, 1958, 1960a, 196lb, 1962, 1965a; Valkanov, 1957; Furnestin, 1959, 1960; Holmquist, 1959b; Bainbridge, 1960; Reys, 1960; Clarke, 1961a; Hoenigman, 1961, 1963, 1968; Pillai, 1961, 1963c, 1964, 1965, 1973; Wickstead, 1961; Salvat, 1962; Hodge, 1963b; Kuhlmorgen-Hille, 1963; Balasubrahmanyan, 1964; Brown, 1964, 1971; Costa, 1964; Ii, 1964a, b ; Sando, 1964; Engel and Tan, 1965; Hopkins, 1965, 1966; Macquart-Moulin, 1965, 1973b, 1977a, c; Zatkutskiy, 1965, 1970; Dediu, 1966; Ariani, 1967; Fishelson and Loya, 1968; O’Riordan, 1969; Riedl and Ozretic, 1969;Brattegard, 1970a; Champelbert and Macquart Moulin, 1970; Fraser, 1970; Arntz, 1971, 1978a, b ; Baan and Holthuis, 1971; Day et al., 1971; Field, 1971; Mauchline, 1971e,f,g, 1972,, 1973b, c; Moran and Fishelson, 1971; Brown and Talbot, 1972; Kuhl and Mann, 1972; Lagardbre, 1972, 1977a, b ; Moran, 1972; Ghirardelli, 1973; Hesthagen, 1973; Shyamasundari, 1973; Vader, 1973; Bacescu and Schiecke, 1974; Hamond, 1974; Hatzakis, 1974, 1977; Gauld and Raju, 1975; Ariani and Spagnuola, 1976; Balasubrahmanyan and Prince, 1976; Boysen, 1976; Nath and Pillai, 1976; Bacescu and Muradian, 1977; Beaudouin, 1977; Boyden et al., 1977; Makings, 1977; Mauchline and Murano, 1977; Warwick and Davies, 1977; Wooldridge, 1977; Hesthagen and Gjermundsen, 1979; McLachlan et al., 1979; Moiler, 1979.

Gibberythrops Ii, 1964a; Pillai, 1964, 1965; Murano, 1970c; Mauchline and Murano, 1977.

350

THE BIOLOGY OF MYSIDS

Gnathophausia Norman, 1880a ; Alcock and Anderson, 1894; CoutiBre, 1904; Stebbing, 1908, 1917; Ostenfeld and Wesenberg-Lund, 1909; Zimmer, 1912; Daniel, 1933; Nouvel, 1941; Boschma, 1949, 1959; Fraser, 1950, 1961; Banner, 1954a; Raw, 1956; Bruun, 1957; Birstein and Tchindonova, 1958, 1962; Fisher and Kon, 1959; Tchindonova, 1959; Brunel, 1960; Zenkevich and Birstein, 1960; Clarke, 1961s; Aron, 1962; Gordon, 1964; Grindley and Penrith, 1965; Pequegnat, 1965; 0. S. Tattersall, 1965b; Holthuis and Sivertsen, 1967; Lewis, 1967; Childress, 1968, i971a, b, 1975; O’Riordan, 1969; Raymont et al., 1969; Taniguchi, 1969, 1976; Murano, 1970c; Vinogradov, 1970; Vinogradov et al., 1970; Lee et al., 1971; Mauchline, 1972, 1973b; Lee and Hirota, 1973; Morris and Sargent, 1973; Vader, 1973; Childress and Nygaard, 1974; Pearcy and Ambler, 1974; Fuzessery and Childress, 1975; Belman and Childress, 1976; Bordovskiy et al., 1976; Casanova, 1977; Lagardhre, 1977a, b ; Mauchline and Murano, 1977; Pearcy et al., 1977; Freel, 1978; MacDonald and Gilchrist, 1978; Mickel and Childress, 1978; Quetin et al., 1978.

Gymnerythrops Pillai, 1973; Mauchline and Murano, 1977.

Hansenomysis Holmquist, 1957b;Birstein and Tchindonova, 1958,1970; Gordon, 1960; Clarke, 1961a; 0. S. Tattersall, 1961a, 1965b; Bacescu, 1967, 1971b; O’Riordan, 1969; Mauchline, 1972; Mauchline and Murano, 1977.

Haplostylus Bacescu, 1973c, 1975; Hatzakis, 1977; Mauchline and Murano, 1977.

Hemimysis Walker, 1888a; Scott, 1896a, 1897; Ostenfeld, 1906; Ostenfeld and Wesenberg-Lund, 1909; Bacescu, 1938, 1954; Liao, 1951; FOX,1952; Fisher et al., 1955; Dahl, 1956; Grainger, 1956; Anonymous, 1957; Valkanov, 1957; Fisher and Kon, 1959; Hoenigman, 1960; Zhuravel, 1960b; Holmquist, 1962; Ledoyer, 1963, 1965, 1966; Gordon, 1964; Dediu, 1966; Macquart-Moulin and Patriti, 1966; Davis, 1968; O’Riordan, 1969; Zatkutskiy, 1970; Mauchline, 1971e, f, g, 1972, 1973b, c; Hesthagen, 1973; Hamond, 1974; Cinelli et al., 1977; LagardBre, 1977b; Makings, 1977; Mauchline and Murano, 1977; Skjoldal and Bamstedt, 1977;Wittmann, 1978;Mordukhai-Boltovskoi, 1979.

APPENDIX II

351

Hemkirielh Bdasubrahmanyan, 1964; Ii, 1964a; Pillai, 1964, 1965, 1973; 0. S. Tattersall, 1965a; Taniguchi, 1976; Mauchline and Murano, 1977.

Heteroerythrops Murano, 1966b; Mauchline and Murano, 1977.

Heteromysis Scott, 1889; Walker, 1898a, b ; Delamare Deboutteville, 1950; Lochhead, 1950; Jones, 1954; Le Sueur, 1954; Anonymous, 1957; Belloc, 1959; Brunel, 1960; Pillai, 1961, 1964, 1965; 0. S. Tattersall, 1961b, 1962, 1967; Richards, 1963; Ii, 1964a; Nouvel, 1964b; Richards and Riley, 1967; Becescu, 1968b, 1970b, 1975, 1976; Brattegard, 1969, 1970b, 1973, 1974a, b, 1975; O’Riordan, 1969; Mauchline, 19718, f, g, 1973b; Wigley and Burns, 1971; Lagardisre, 1972, 1977b; Williams, 1972; Hamond, 1974; Bacescu and Muradian, 1977; Camp et al., 1977; Makings, 1977; Mauchline and Murano, 1977.

Heteromysoides Bacescu, 1968b; Mauchline and Murano, 1977.

Holmsiella Banner, 1954s; Biretein and Tchindonova, 1958; Ii, 1964a; Murano, 1970b, c, 1976; Ikeda, 1977; Mauchline and Murano, 1977.

Hyperamblyops Birstein and Tchindonova, 1958; Vinogradov, 1962; O’Riordan, 1969; Zharkova, 1970; Mauchline, 1972; Murano, 1975; Mauchline and Murano, 1977.

Hypererythrops Ii, 1964a; Pillai, 1964, 1965; 0.S. Tattersall, 1965a; Ikeda, 1970; Murano, 1970b, c; Wigley and Burns, 1971; Nouvel, 1974; Nouvel and Lagardhre, 1976; Lagardhre, 1977a, b; Mauchline and Murano, 1977.

H yperiimysis Nouvel, 1966b; Bacescu, 1975; Mauchline and Murano, 1977.

Idiompis Pillai, 1964, 1965; Bacescu, 1973a; Mauchline and Murano, 1977; Murano, 1978.

352

TEE BIOLOGY OF MYSIDS

Iiella Bacescu, 1968d; Mauchline and Murano, 1977.

Iimysis Nouvel, 1966a; Mauchline and Murano, 1977.

Indomysis Gordon, 1960; Clarke, 1961a; Pillai, 1964,1965; Mauchline and Murano, 1977.

Inusitatomysis Banner, 1954a; Ii, 1964a; Mauchline and Murano, 1977.

Kainomatomysis Bacescu, 1973d, 1975; Mauchline and Murano, 1977.

Katamysis Bacescu, 1938, 1954; Holmquist, 1962; Dediu, 1966; Mauchline, 197313; Mauchline and Murano, 1977 ; Mordukhai-Boltovskoi, 1979.

Katerythrops Zimmer, 1912; Banner, 1954a; Fraser, 1961; Ii, 1964a; O’Riordan, 1969; Murano, 1976; Mauchline, 1977a; Mauchline and Murano, 1977; Panampunnayil, 1977.

Leptom ysis Scott, 1889, 1897, 1900a; Walker, 1898b; Ostenfeld, 1906, 1916; Ostenfeld and Wesenberg-Lund, 1909, Zimmer, 1912; Colosi, 1923; Hardy, 1924; Vannini, 1930; Moore, 1937; Bacescu, 1938, 1954, 1966; Santa and Bacescu,l942; Liao, 1951; Nouvel, 1951a, 1973c; Fisher et al., 1953; Jones, 1954; Le Sueur, 1954; Colman and Segrove, 1955; Nouvel and Hoenigman, 1955; Anonymous, 1957; Valkanov, 1967; Hoenigman, 1958a, b, 1960, 1963, 1968; Boschma, 1959; Fisher and Kon, 1959; Furnestin, 1959, 1960; Massonet, 1960; Reys, 1960; Ben’ko, 1962; Pillai, 1964, 1965, 1973; Rice, 1964; Siudzinski, 1964; Macquart-Moulin, 1965, 1970, 19738, b ; Vives, 1965, 1968; Ledoyer, 1966; Raymont and Linford, 1966; Ariani, 1967; Seguin, 1968; Mauchline, 1969a, 1971e, f, g, 1972, 1973a, b, c; O’Riordan, 1969; Champelbert and Macquai-t-Moulin, 1970; Lagardkre, 1972, 197%; Ghirardelli, 1973; Hesthagen, 1973; Shyamasundari, 1973; Vader, 1973; Hamond, 1974; Hatzakis, 1974; Wing, 1975; Beaudouin, 1977;

APPENDIX 11

353

Makings, 1977; Maucuine and Murano, 1977; Wittmann, 1977, 1978; Lucu, 1978.

Limnornysis Bacescu, 1936a, 1938, 1954; Woparovich, 1954; Valkanov, 1957; Zhuravel, 1959, 1960a, 1972; Holmquist, 1962; Dediu, 1966; Ivanov, 1972; Mauchline, 1973b; Weish and Turkay, 1975; Mauchline and Murano, 1977; Boroditch, 1978; Mordukhai-Boltovskoi, 1979.

Longithorax Banner, 1954a; Birstein and Tchindonova, 1958; Belloc, 1959; 0. S. Tattersall, 1961b; Vinogradov, 1962; Pillai, 1973; Murano, 1976; Mauchline and Murano, 1977.

Lophogaster Walker, 189810; CoutiBre, 1904; Ostenfeld and Wesenberg-Lund, 1909; Zimmer, 1912; Pawsey and Davis, 1924; Broch, 1927; Daniel, 1933; Banner, 1954a; Raw, 1955; Furnestin, 1959; Reys, 1960; 0. S. Tattersall, 1960b, 196lb; Bacescu and Mayer, 1961; Clarke, 1961a; Curl, 1962; Hoenigman, 1963, 1968; Gordon, 1964; Elofsson, 1965; O’Riordan, 1969; Casanova, 1970, 1977; Elofsson and Dahl, 1970; Fraser, 1970; Murano, 1970a; Franqueville, 1971; Wiborg, 1971; Lagardhre, 1972, 1977a, b ; Mauchline, 1972, 197313; Morris, 1972; Macquart-Moulin ,197313; Nouvel, 1973c, 1978; Pillai, 1973; Bacescu, 1976; Bacescu and Muradian, 1977; Mauchline and Murano, 1977. Lycomysis

Colosi, 1916, 1923; Pillai, 1961, 1964, 1965; Ii, 1964a; Mauchline and

Murano, 1977. Mesacanthornysis Nouvel, 1967; Mauchline, 1973b; Bacescu, 1975; Mauchline and Murano, 1977.

Mesopodopsis Henderson, 1885; Scott, 1889, 1900b, 1902b; Ostenfeld, 1906, 1916; Ostenfeld and Wesenberg-Lund, 1909; Mielck, 1911; Zimmer, 1912; Hardy, 1924; Schodduyn, 1926; Percival, 1929; Savage, 1931; Otto and Wielinga, 1933; Bacescu, 1938, 1954; Nouvel, 1951a, 1954, 1957a, 1973c, 1978; Le Sueur, 1954; Colman and Segrove, 1955; Anonymous, 1957; Caspers, 1957; 0. S. Tattersall, 1957, 1960a, 1965a; Valkanov, 1957; Holmquist, 1959b; Bainbridge, 1960; Furnestin, 1960; Pillai,

354

THE BIOLOGY OF MYSIDS

1961, 1964, 1965; Wickstead, 1961; Ben’ko, 1962; Mankowski, 1962; Linford, 1965; Macquart-Moulin, 1965, 1970; Zatkutskiy, 1965, 1970; Dediu, 1966; Ariani, 1967; Davis, 1968; Hoenignian, 1968; Nekrasova and Rakitina, 1968; Vives, 1968; Baan and Holthuis, 1971; Mauchline 1971e, g, 1972, 1973b; Kaliyamurthy, 1972; Kuhl c;iLdMann, 1972; Ghirardelli, 1973; Hesthagen, 1973; Raymont et al., 1973; Hamond, 1974; Greve, 1975; Krylova et al., 1975; Silas and Pillai, 1975.;Boysen, 1976; Boyden et al., 1977; Makings, 1977; Mauchline andMurano, 1977; Win, 1977 ; Wittmann, 1977 ; Wooldridge, 1977 ; Whitfield and Blaber, 1978.

Metamblyops Ii, 1964a; O’Riordan, 1969; Mauchline and Murano, 1977. Metam ysidopsis Clarke, 1961a; Costa, 1964; Fleminger and Clutter, 1966; Hopkins, 1965, 1966; Clutter, 1967; Bacescu, 1968c, 1969; Fager and Clutter, 1968; Brattegard, 1970a, b, 1973, 1974a, b, 1975; Silva, 1970b, 1977; Clutter and Theilacker, 1971; Day et al., 1971; Almeida Prado, 1972, 1973, 1974; Williams, 1972; Mauchline, 1973b; Quintero and Roa, 1973, 1977; Price, 1975; Mauchline and Murano, 1977. Meterythrops Ostenfeld and Wesenberg-Lund, 1909; Stephensen, 1943; Banner, 1954a; Birstein and Tchindonova, 1958, 1962; Tchindonova, 1959; Brunel, 1960, 1961, 1970; Ii, 1964a; Pillai, 1964, 1965; O’Riordan, 1969; Poirier, 1969; Taniguchi, 1969; Murano, 1970c, 1977a; Wigley and Burns, 1971; Wing, 1975; Mauchline and Murano, 1977; Shih and Laubitz, 1978. Michthyops Stephensen, 1943; Geiger, 1969; O’Riordan, 1969; LagardAre, 19’77%b ; Mauchline and Murano, 1977. Mysideis Ostenfeld and Wesenberg-Lund, 1909; Zimmer, 1912; Pawsey and Davis, 1924; Reys, 1960; O’Riordan, 1969; LagardAre, 1972, 1977a, b ; Mauchline and Murano, 1977. Mysidella Ostenfeld and Wesenberg-Lund, 1909; Banner, 1954a; Raw, 1955; Ii, 1964a; O’Riordan, 1969; Murano, 1970b, c; Brattegard, 1973; LagardAre, 1977b; Mauchline and Murano, 1977.

APPENDIX 11

355

Mysidetes Rustad, 1930; Holmquist, 1957b; 0. S. Tattersall, 1961a, 1965b; Bacescu, 1967; Ledoyer, 1969; O’Riordan, 1969; Mauchline, 197313; Lagardhe, 1977a, b ; Mauchline and Murano, 1977.

Myaidium Bainbridge and Waterman, 1957, 1958; Jander and Waterman, 1960; Steven, 1961;Jander, 1962; Costa, 1964; Randall eta,?.,1964; Goodbody 1965; Davis, 1966, 1968; Eguchi and Waterman, 1966, 1973; Emery, 1968; Brattegard, 1969, 1970b, 1973, 1974a, b, 1975; Almeida Prado, 1972, 1974; Mauchline, 1972, 1973b; Camp et al., 1977; Mauchline and Murano, 1977; Quintero and Roa, 1977; Silva, 1977.

Mysidopsis Norman, 1865; Scott, 1889, 1891b, 1896a, 1897, 1900a, 1902a; Walker, 1898b; Oste’nfeld, 1906, 1916; Ostenfeld and Wesenberg-Lund, 1909; Zimmer, 1912; Moore, 1937; Liao, 1951; Le Sueur, 1954; Anonymous, 1957; Bossanyi, 1957; Bowman, 1957, 1964; Holmquist, 195713; Darnell, 1958, 1961; Hopkins, 1958, 1965, 1966; 0. S. Tattersall, 1958, 196lb, 1962, 196510, 1969; Fisher and Kon, 1959; Furnestin, 1959; Reys, 1960; Grice and Hart, 1962; Ii, 1964a, b ; Nouvel, 1964a, 1973d; Pillai, 1964, 1965, 1967; Sando, 1964; Engel and Tan, 1965; MacquartMoulin, 1965; Ledoyer, 1966; Ariani, 1967; Clutter, 1967; Bacescu, 1968c, 1969, 1975; Hoenigman, 1968; Mauchline, 1968, 1970b, 1971f, g, 1972, 1973a, b, c, 1977a; Brattegard, 1969, 1970a, 1973, 1974a, b, 1975; Molenock, 1969; O’Riordan, 1969; Champelbert and Macquart-Moulin, 1970; Murano, 1970b, c; Baan and Holthuis, 1971; Day et al., 1971; Field, 1971; Wigley and Burns, 1971; Almeida Prado, 1972,1973,1974, Williams, 1972; Bacescu and Vasilescu, 1973; Ghirardelli, 1973; Hesthagen, 1973; Marsh, 1973; Hamond, 1974; Ogle and Price, 1976; Hallberg, 1977; Lagardhre, 1977b; Makings, 1977; Mauchline and Murano, 1977; Schimmel et al., 1977; Silva, 1977; Christensen, 1978; Lawler and Shepard, 1978; Nimmo et aZ., 1978; Overstreet and Heard, 1978; Price, 1978; Price and Vodopich, 1979.

Mysimenzies Bacescu, 1971a; Mauchline and Murano, 1977.

Mysis Scott, 1899; Kane, 1904; Ostenfeld, 1906, 1916; Ostenfeld and Wesenberg-Lund, 1909; Walton, 1922; Stephensen, 1933, 1943;

356

THE BIOLOUY OF MYSIDS

Langford, 1938a, b ; Rawson, 1942, 1960; Larkin, 1948; Mohr, 1953; Banner, 1954a; Dahl, 1956; Holmquist,1957, 1958a, b, 1959a, b,1962, 1963a, b, 1966, 1973; Segerstrble, 1957, 1962; Jensen, 1958; McLaren, 1958; Beeton, 1959, 1960, 1969; Lindquist, 1959; Ricker, 1959; Brunel, 1960, 1961, 1970; Hynes et al., 1960; Wells, 1960; Clarke, 1961a; Grainger, 1962 ;Lettevall, 1962 ;Prefontaine and Brunel, 1962 ; Shurin, 1962; Zmudzinski, 1962; Richards, 1963; Wells and Beeton, 1963; Haahtela, 1964; Johnson, 1964; Sparrow et al., 1964; Siudzinski, 1964; Dryer et al., 1965; Elofsson, 1965; Furst, 1965, 1967; Grimas and Nilsson, 1965; Linn and Frantz, 1965; Jolicoeur and Brunel, 1966; McNaught and Hasler, 1966; Schumacher, 1966; Stringer, 1967; Vinogradova and Kandiuk, 1967; Vinogradova and Petkevich, 1967 ; Haefner, 1968, 1969; McNaught, 1968; Morswell and Norden, 1968; Powers and Robertson, 1968; Ackefors, 1969; Fiirst and Nyman, 1969; Geiger, 1969; Hogstad, 1969; Kulakovskii, 1969, 1971, 1974; Nyman and Westin, 1969; Poirier, 1969; Schneider et al., 1969; Tracy and Vallentyne, 1969; Uzars, 1969, 1972; Brownell, 1970; Elofsson and Dahl, 1970; Just, 1970; Khlebovich et al., 1970; Reger et al., 1970; Smith, 1970; Swain et al., 1970; Westin, 1970; Zyblut, 1970; Amtz, 1971, 1978a; Wigley and Burns, 1971; Beeton and Edmondson, 1972; Kinney, 1972; Lasenby and Langford, 1972, 1973; Northcote, 1972, 1973; Reynolds and DeGraeve, 1972; Saether and McLean, 1972; Sarkka, 1972, 1976; Teraguchi et al., 1972, 1975; Thommes et al., 1972; Tyler, 1972, 1973; Zelikman and Golovkin, 1972; Bagge and Ilus, 1973; Barker, 1973; Dormaar and Corey, 1973, 1978; Eguchi and Waterman, 1973; Healey and Woodall, 1973; Hesthagen, 1973; Mauchline, 1973b; Nikolaev et al., 1973; Sergeant, 1973; Carpenter et al., 1974; Judkins and Wright, 1974; Prygunkova, 1974; Shrivastava, 1974; Watson, 1974; Yaguchi et al., 1974; Amaratunga and Corey, 1975; Dadswell, 1975; Fain-Maurel et al., 1975a; Johnson, 1975; Marshall et al., 1975; Nuorteva and Hasanen, 1975; Richards et al., 1975; Savolainen, 1975; Wiens et al., 1975; Wing, 1975; Biesinger et al., 1976; Foulds and Roff, 1976; Salonen et al., 1976; Vinogradov, 1976; Bowman, 1977a; Boysen, 1976; Mauchline and Murano, 1977; Wing and Barr, 1977; Borgmann, 1978; Bowers and Grossnickle, 1978; Foulds and Mann, 1978; Hakala, 1978; Morgan and Beeton, 1978; Shih and Laubitz, 1978; Grossnickle and Beeton, 1979; Grossnickle and Morgan, 1979; Pezzack and Corey, 1979. Nanomysis

Nouvel, 1957a; 0. S. Tattersall, 1960a; Wickstead, 1961; Ii, 1964a; Mauchline and Murano, 1977.

APPENDIX II

357

Neohteromysis Bacescu, 1976; Maychline and Mkano, 1977.

Neornysis Walker, 1888a; Scott, 1891a, 1894, 1896a, 1897, 19OOa, 1902b; Ostenfeld, 1906; Ostenfeld and Wesenberg-Lund, 1909; Mielck, 1911; McIntosh, 1919; W. Tattersall, 1919; Percival, 1929; Otto and Wielinga, 1933; Shelford, 1935; Rees, 1939; Havinga, 1941; Redeke, 1941; Vos, 1941; Santia and Bacescu, 1942; Whiteley, 1948; Ishikawa and Oshima, 1951; Banner, 1954a; Le Sueur, 1954; Light, 1954; Suomalainen, 1954; Fisher et al., 1955; Dahl, 1956; Grainger, 1956; Anonymous, 1957; Holmquist, 1957a, b, 1959b, 1973; Hulbert, 1957; Hopkins, 1958, 1965; Jepsen, 1958; Fisher and Kon, 1959; Furnestin, 1959; Lindquist, 1959; Brunel, 1960; Hynes et a,?.,1960; Raymont and Krishnaswamy, 1960, 1968; Clarke, 1961a; Raymont and Conover, 1961; Wiktor, 1961; Yamada, 1961, 1964; Cronin et al., 1962; Fukai et al., 1962; Grant, 1962; Grice and Hart, 1962; Herman, 1962, 1963; Prefontaine and Brunel, 1962; Price, 1962; Zmudzinski, 1962; Heubach et al., 1963; Murano, 1963a, b, 1964a, b, 1966c, d, e,; Richards 1963; Haahtela, 1964; Hirano et al., 1964a, b ; Ii, 1964a; Raymont et al., 1964, 1966, 1967, 1968, 1973; Siudzinski, 1964; Elofsson, 1965; Engel and Tan, 1965; Linford, 1965; 0.S. Tattersall, 1965b; Wing, 1965,1975; Gansale, 1966; Painter, 1966; Radtke, 1966; Raymont and Linford, 1966; Sasaki, 1966a, b, c; Stevens, 1966a, b ; Turner, 1966a, b ; Turner and Heubach, 1966; Clutter, 1967; Hwrtel and Osterberg, 1967; Miller, 1967, 1970; MUUS, 1967; Richards and Riley, 1967; Thomas, 1967; Tundisi and Krishnaswamy, 1967; Herman et al., 1968; Jansson and Kiillander, 1968; Schlieper, 1968; Seguin, 1968; Adams, 1969; Fujika et al., 1969; Haefner, 1969; Heubach, 1969; Jawed, 1969, 1973; Nyman and Westin, 1969; O’Riordan, 1969; Elofsson and Dahl, 1970; Markle and Grant, 1970; Schaefer, 1970; Barnes et d., 1971; Berrill, 1971; Burton and Massie, 1971; Chin, 1971, 1972, 1974, 1976; Friedl, 1971a, b ; Hair, 1971; Lee and Chin, 1971; McLusky and Heard, 1971; Mauchline, 1971b, f, g, 1972, 1973a, b ; Morris, 1971, 1973; Okada and Taniguchi, 1971; Shushkina et al., 1971; Srinivasagam et al., 1971; Teshima and Kanazawa, 1971; Wigley and Burns, 1971; Kuhl and Mann, 1972; Sage and Herman, 1972; Shushkina, 1972; Sikora et al., 1972; Swartz, 1972; Werner, 1972; Williams, 1972; Elofsson and Hallberg, 1973; Marsh, 1973; Morris et a,!., 1973; Nouvel, 1 9 7 3 ~ ; Shyamasundari, 1973; Vader, 1973; Boesch and Diaz, 1974; Hamond, 1974; Jacobs and Grant, 1974; Kasaoka, 1974; Simmons et al., 1974; Williams et al., 1974; Zelickman, 1974; Hirano, 1975; Kost and Knight,

358

THE BIOLOGY O F MYSIDS

1975; Klyashtorin and Kuz’micheva, 1975; Nuorteva and Hiisiinen, 1975; Pechen-Finenko and Pavlovskaya, 1975; Simmons and Knight, 1975; Stickney and Knowles, 1975; Stickney et al., 1975; Wiens et al., 1975; Boesch et a1.,1976; Boysen, 1976; Kislalioglu and Gibson, 1976a, b ; Kravitz et al., 1976; McEachran et al., 1976; Maurer et aZ., 1976; Moore, 1976; Moore and Moore, 1976; Beaudouin, 1977; Bowman, 1977a; Boyden et al., 1977; Hallberg, 1977; Klyashtorin and Musayeva, 1977; Makings, 1977; Mauchline and Murano, 1977; Morris et al., 1977; Pasternak, 1977; Ristich et al., 1977; Vlasblom and Elgershuizen, 1977; Win, 1977; Wing and Barr, 1977; Wittrnann, 1977; Yamamoto, 1977; Eggers et al., 1978; Grabe, 1978; Omori, 1978; Moore et aZ., 1979; Parker, 1979; Parker and West, 1979; Pezzack and Corey, 1979.

Nipponerythrops Mauchline and Murano, 1977; Murano, 1977a. Nouvelia Bacescu and Vasilescu, 1973; Mauchline and Murano, 1977. Paracanthomysis Ii, 1964a; Ogawa, 1964; Lee and Chin, 1976; Mauchline and Murano, 1977. Paralophogaster Clarke, 1961a; Pillai, 1973; Mauchline and Murano, 1977. Paramblyops Ostenfeld and Wesenberg-Lund, 1909; 0. S. Tattersall, 1965b; O’Riordan, 1969; Birstein and Tchindonova, 1970; LagardBre, 1972, 1977a, b ; Nouvel and Lagardike, 1976; Mauchline and Murano, 1977. Paramysis Herdman, 1920; Bacescu, 1936a, 1938, 1954; Moore, 1937; Santa and Bacescu, 1942; Liao, 1951; Gabe, 1952; Fisher et al., 1953; Labat, 1954,1957; Le Sueur, 1954; Anonymous, 1957; Caspers, 1957; Valkanov 1957; Fisher and Kon, 1959; Furnestin, 1959; Zhuravel, 1959, 1960a, 1972; Massonet, 1960; Hoenigman, 1961; Ben’ko, 1962; Holmquist, 1962; Salvat, 1962; Juchault, 1963; Meusy, 1963; Nouvel, 1964c, 1973c; Rice,4964; Macquart-Moulin, 1965; Nouvel and Panouse, 1965; Zatkutskiy, 1965, 1970; Charniaux-Cotton et al., 1966; Dediu, 1966; TskhomeKke and Sergeeva, 1966; Ariani, 1967; Vinogradova and Kandiuk,7967 ; Vinogradova and Petkevich, 1967 ; Nekrasova and

APPENDIX II

359

Rakitina, 1968; O’Riordan, 1969; Boroditch and Havlena, 1971, 1973; Mauchline, 1971a, f,’g, 1972, 1973a, b ; Belyayeva et al., 1972; Ivanov, 1972; Kuhl and Mann, 1972; Ghirardelli, 1973; Miroshnichenko and Vovk, 1973; Hamond, 1974; Krylova et al., 1975; Savchuk, 1975; Bacescu and Muradian, 1977; Makings, 1977, 1978; Mauchline and Murano, 1977; Boroditch, 1978; Mordukhai-Boltovskoi, 1979.

Paranchialina Taniguchi, 1976; Mauchline and Murano, 1977. Parapseudomma Nouvel and Lagardhe, 1976; Lagardhe, 1977a, b ; Mauchline and Murano, 1977. Parastilmysis Enomoto, 1964; Ii, 1964a; Murano, 1970b, c; Mauchline and Murano, 1977.

Parerythrops Ostenfeld and Wesenberg-Lund, 1909; Stephensen, 1943; Birstein and Tchindonova, 1959; O’Riordan, 1969; Brattegard and Vader, 1972; Nouvel and LagardGre, 1976; Lagardhre, 1977b; Mauchline and Murano, 1977. Petalophthalmzcs Alcock and Anderson, 1894; Dakin and Colefax, 1940; Birstein and Tchindonova, 1958; Tchindonova, 1959; Gordon, 1960; Clarke, 1961a; Vinogradov, 1962; Pillai, 1965, 1968, 1973; 0. S. Tattersall, 1968; Murano, 1970c; Zharkova, 1970; Mauchline, 1972; Mauchline and Murano, 1977. Pleurerythrops Ii, 1964a; 0. S. Tattersall, 1965%;Murano, 1970b; Mauchline and Murano, 1977; Panampunnayil, 1977. Praunus Norman, 1865; Leslie and Herdman, 1881; Walker, 1886; Scott, 1889, 1896a, 1897, 1900a, 1902b; Herdman, 1890; Meek, 1901, 1903; Ostenfeld, 1906; Ostenfeld and Wesenberg-Lund, 1909; McIntosh, 1919; Anonymous, 1920; Scott, 1922; Broch, 1927; Daniel, 1928, 1931, 1932; Percival, 1929; Lonnberg, 1931a, 1934; Otto and Wielinga, 1933; Moore, 1937; Sivertsen, 1938; Beanland, 1940; Santa and Bacescu,

360

THE BIOLOQY O F MYSIDS

1942; Nouvel, 1945, 1964c, 19730; Debaisieux, 1947, 1949; Liao, 1961; Smidt, 1951; Gabe, '1952; Le Sueur, 1954; Alexandrowicz, 1955; Colman and Segrove, 1955; Mapat, 1955, 1965a, b, 1962; Dahl, 1956; Anonymous, 1957; Holmquist, 1957a, 1959b; Segerstrhle, 1957; Jensen, 1958; Lindquist, 1959; Nicol, 1959; Massonet, 1960; Labat, 1961; Raymont, 1961; Rice, 1961, 1964; Molander, 1962; Salvat, 1962; Zmudzinski, 1962; Juchault, 1963; Wigley, 1963; Siudzinsky, 1964; Elofsson, 1965; Linford, 1965; Mauchline, 1965, 1971c, f, g, 1972, 1973a, b ; 1977a; Chamiaux-Cotton et al., 1966; Muus, 1967; Jansson and Kallander, 1968; Seguin, 1968; Haefner, 1968; Nyman and Westin, 1969; O'Riordan, 1969; Elofsson and Dahl, 1970; Baan and Holthuis, 1971; Berrill, 1971; McLusky and Heard, 1971; Wigley and Burns, 1971; Kuhl and Mann, 1972; Saudray, 1972; Elofsson and Hallberg, 1973; Haahtela, 1973; Hesthagen, 1973; Reger and Fain-Maurel, 1973; Hamond, 1974; Fain-Maurel et al., 1975a,b; Greve, 1975; Neil, 1975a, b, c; Nuorteva and H&s&nen,1975; Austin, 1976; Boysen, 1976; Moore, 1976; Moore and Moore, 1976; Boyden et al., 1977; Hallberg, 1977; Laverack et al., 1977; Makings, 1977, 1978; Mauchline and Murano, 1977 ; Vlasblom, and Elgershuizen, 1977 ; Parker, 1979; Pezzack and Corey, 1979.

Prionom ysis Ii, 1964a; Pillai, 1964, 1965; Mauchline and Murano, 1977. Promysis 0. S . Tattersall, 1960a, 1965a; Wickstead, 1961; Costa, 1964; Ii, 1964a; Pillai, 1964, 1965, 1973; Hopkins, 1966; Day et al., 1971; Wigley and Burns, 1971; Almeida Prado, 1972, 1973, 1974; Williams, 1972; Brattegard, 1973, 1974a; Mauchline and Murano, 1977; Silva, 1977. Proneomysis Banner, 1954a; Ii, 1964a, b ; Sando, 1964; Kikuchi, 1967; Mauchline and Murano, 1977; Murano, 197713. Pseudamblyops Ii, 1964a; Mauchline and Murano, 1977. Pseudanchialina Nouvel, 1944; 0. S. Tattersall, 1960a, 1965a; Wickstead, 1961; Balasubrahmanyan, 1964; Ii, 1964a; Pillai, 1963, 1965, 1973; Shyamasundari, 1973; Bacescu, 1975; Taniguchi, 1976; Mauchline and Murano, 1977; Sale et al., 1978.

APPENDIX 11

361

Pseuderythrops Nouvel, 1944; Pillai, 1964, 1965, 1973; Mauchline and Murano, 1977. Pseudochalaraspidum Birstein and Tchindonova, 1962; Mauchline and Murano, 1977. Pseudomma Ostenfeld, 1906; Ostenfeld and Wesenberg-Lund, 1909 ; Rustad, 1930; Stephensen, 1943; Banner, 1954a; Klawe, 1955; Holmquist, 195713; Birstein and Tchindonova, 1958; Brunel, 1960, 1961, 1970; Reys, 1960; 0.S. Tattersall, 1961a, 1965b; Ii, 1964a; Murano, 1966a, 1970b, c, 1974b; Geiger, 1969; O’Riordan, 1969; Poirier, 1969; Mauchline, 1971e,f,g, 1972, 1973b, c ; Wigley and Burns, 1971; Nouvel and Lagardbre, 1976 ; Elofsson and Hallberg, 1977 ; Lagerdhre, 1977b; Mauchline and Murano, 1977; Hopkins and Gulliksen, 1978. Pseudomysidetes Mauchline and Murano, 1977 ; Panampunnayil, 1977. Pseudomysis Ii, 1964a; Murano, 1970c; Mauchline and Murano, 1977. Pseudoxomysis Nouvel, 1973a; Mauchline and Murano, 1977. Pteromysis Ii, 1964a; Murano, 1970c, 1976; Mauchline and Murano, 1977. Rhopalophthalmus Zimmer, 1912; Dakin and Colefax, 1940; 0. S. Tattersall, 1957, 1958, 1960a, 1961b, 1965a; Furnestin, 1959; Bainbridge, 1960; Nouvel, 1960; Pillai, 1961, 1964, 1965, 1973; Wickstead, 1961; Hodge, 1963a,b; Balasubrahmanyan, 1964; Ii, 1964a ;Mauchline, 197313;Shyamasundari, 1973; Mauchline and Murano, 1977; Wooldridge, 1977. Schistomysis Norman, 1865; Walker, 1886, 1898b; Herdman, 1890; Scott, 1897, 1902b; Meek, 1901, 1903, 1908, 1909; Ostenfeld, 1906, 1916; Ostenfeld and Wesenberg-Lund, 1909; Mielck, 1911; Scott, l”922; Hardy, 1924; Percival, 1929; Savage, 1931; Moore, 1937; Rees, 1939; Santa and Bacescu, 1942; Liao, 1951; Nouvel, 1951a, 1973c; Jones, 1954; Le Sueur, 1954; Colman and Segrove, 1955; Anonymous, 1957; Bossanyi,

362

THE BIOLOGY OF MYSIDS

1957; Rice, 1961, 1964; Elofsson, 1965; Nagabhushanan, 1965; Almeida Prado, 1966; Mauchline, 1967, 1970a, 1971d, f, g, 1972, 1973a, b, c, 1977a; Edwards and Steele, 1968; O’Riordan, 1969; Baan and Holthuis, 1971; Brattegard and Vader, 1972; Kuhl and Mann, 1972; Shotter, 1972; Hesthagen, 1973; Hamond,1974; Boysen, 1976; Beaudouin, 1977; Boyden et al., 1977; Makings, 1977, 1978; Mauchline and Murano, 1977.

Scolamblyops Murano, 1974a; Mauchline and Murano, 1977.

Xiriella Kinahan, 1859; Leslie and Herdman, 1881; Scott, 1887, 1897, 19OOa, 1902a; Walker, 1888b, 1898b; Herdman, 1890, 1920; Holt and Beaumont, 1899; Ostenfeld, 1906, 1916 ; Ostenfeld and Wesenberg-Lund, 1909; McIntosh, 1919; Savage, 1931; Moore, 1937; Bacescu, 1938, 1954, 1973d, 1975; Dakin and Colefax, 1940; Sen& and Bacescu, 1942; Nouvel, 1944, 1951a, 1952, 1957a, 1 9 7 3 ~ ;Liao, 1951; Fox, 1952; Gabe, 1952; Fisher et al., 1953; Banner, 1954a; Le Sueur, 1954; Colman and Segrove,1955; Genovese, 1956, 1957; Anonymous, 1957; Holmquist, 1957a, b, 1959b; Valkanov, 1957; Hoenigman, 1958a, b, 1960, 1963, 1964a, b, 1965, 1968; 0. s. Tattersall, 1958, 1960a, 1961b, 1962, 1965a; Belloc, 1959; Fisher and Kon, 1959; Furnestin, 1959, 1960; Dion andNouvel,1960; Massonet, 1960; Reys, 1960; Bacescu and Mayer, 1961; Clarke, 1961a; Evans, 1961; Pillai, 1961, 1964, 1965, 1973; Rice, 1961, 1964; Wickstead, 1961; Ben’ko, 1962; Birstein and Tchindonova, 1962 ; Balasubrahmanyan, 1964;Costa, 1964;Ii, 1964a, b ; Sando, 1964; Ledoyer, 1965, 1966, 1970; Macquart-Moulin, 1965, 1970, 1972, 1973b; Zatkutskiy, 1965, 1970; Bellan-Santini, 1966; Ariani, 1967 ; Kikuchi, 1967; Omori, 1969a; O’Riordan, 1969; Brattegard, 1970a, b, 1973, 1974a, b, 1975; Champelbert and Macquart-Moulin, 1970; Baan and Holthuis, 1971; Mauchline, 1971e, f, g, 1972, 1973b, c; Wiborg, 1971; Almeida Prado, 1972, 1974; Ghirardelli, 1973; Shyamasundari, 1973; Vader, 1973; Hamond, 1974; Hatzakis, 1974; Silva, 1974, 1977; Ariani and Spagnuola,l976; Austin, 1976; Hobson and Chess, 1976; Taniguchi, 1976; Boyden et al., 1977; Hallberg, 1977; Lagardhre, 1977b; Makings, 1977; Mauchline and Murano, 1977; Moore and Sander, 1977; Pratap et al., 1977; Quintero and Roa, 1977; Wittmann, 1977, 1978.

Spelaeomysis Gordon, 1960; Clarke, 1961b; Pillai and Mariamma, 1964; Pillai, 1965; Riedl and Ozretic, 1969; Bacescu and Orghidan, 1971; Nath, 1972,

APPENDIX II

363

1973, 1974; Nath and Pillai, 1971; Ingle, 1972; Nath et al., 1972; Bowman, 1973, 1977%; Mauchline, 1973b; Pesce, 1975a; Mauchline and Murano, 1977.

Stilomysis Stephensen, 1943; Brunel, 1960, 1961, 1970; Ii, 1964a; Vader, 1973; Wing, 1975; Mauchline and Murano, 1977; Hopkins and Gulliksen, 1978. St ygiomysis Caroli, 1937; Gordon, 1958, 1960; Clarke, 1961a; R e i d and Ozretic, 1969; Nath and Pillai, 1971; Pesce, 1975b; Bowman, 1976; Mauchline and Murano, 1977. Synerythrops Birstein and Tchindonova, 1958; Ii, 1964a; Pillai, 1964, 1965; Murano, 1975; Mauchline and Murano, 1977. Taphromysis Bacescu, 1961; Hopkins, 1966; Kohout and Kolipinsky, 1967; Brattegard, 1969; Beck, 1977; Bowman, 1977a; Mauchline and Murano, 1977; Johnson and Hopkins, 1978. Tenagomysis Stephensen, 1927; Dakin and Colefax, 1940; Bary, 1956; 0. S. Tattersall, 1957; Belloc, 1959; Hodge, 1964; Ii, 1964a; Nouvel, 1966a; Mauchline, 1973b; Bacescu, 1975; Griffiths, 1976; Mauchline and Murano, 1977. Teraterythrops Ii, 1964a; Murano, 1975; Mauchline and Murano, 1977 Thalassomysis Belloc, 1959; Clarke, 1961a; Pillai, 1964, 1965; Mauchline and Murano, 1977.

Troglomysis Holmquist, 1962 i Mauchline and Murano, 1977.

APPENDIX I11

CLASSIFIED LIST OF LITERATURE FOR GEOGRAPHICAL REGIONS Gordan (1957), in her bibliography of the Mysidacea, classified the literature according to geographical regions. She further divided it into that describing the fauna of coastal as opposed to oceanic regions. The present bibliography does not have this latter sub-division but all papers on both oceanic and coastal species are grouped together for each region. The regions are as shown in Fig. 62 (p. 222), except that the Baltic, Black and Caspian Seas are identified separately as is the literature on freshwater environments.

Freshwater environments Scott, 1891a, 1894; Kane, 1904; W. Tattersall, 1919; Walton, 1922; Langford, 1938a, b ; Rawson, 1942, 1960; Larkin, 1948; Mohr, 1953; Woynarovich, 1954; SegerstrBle, 1957, 1962 ; Holmquist, 1959b, 1962, 1963a, b, 1966, 1973; Mundie, 1959; Ricker, 1959; Zhuravel, 1959, 1960a, b ; Beeton, 1960, 1969; Hynes et al., 1960; Wells, 1960; Lettevall, 1962; Wells and Beeton, 1963; Johnson, 1964, 1975; Sparrow et al., 1964; Dryer, et a1.,1965; Fiirst, 1965, 1967; Grimas and Nilsson, 1965; Linn and Frantz, 1965; McNaught and Hasler, 1966; Schumacher, 1966; Tskhomelidze and Sergeeva, 1966; Stringer, 1967; Morswell and Norden, 1968; Robertson et al., 1968; Schneider et al., 1969; Tracy and Vallentyne, 1969; Brownell, 1970; Smith, 1970, Swain et al., 1970; Zyblut, 1970; Boroditch and Havlena, 1971, 1973; Beeton and Edmondson, 1972; Ivanov, 1972; Kinney, 1972; Lasenby and Langford, 1972, 1973; Northcote, 1972, 1973; Reynolds and DeGraeve, 1972; Saether and McLean, 1972; Siirkka, 1972, 1976; Teraguchi et al., 1972, 1975; Healey and Woodall, 1973; Miroschnichenko and Vovk, 1973; Nikolaev et al., 1973; Carpenter et al., 1974; Shrivastava, 1974; Watson, 1974; Yaguchi et al., 1974; Marshall et al., 1975; Pesce, 1975a; Richards et al., 1975; Weish and Turkay, 1975; Wiens et at.., 1975; Foulds and Roff, 1976; Salonen et al., 1976; Vinogradov, 1976; Beck, 1977;

APPENDIX 111

365

Bowman, 1977a, b ; Borgmann, 1978; Boroditch, 1978; Bowers and Grossnickle, 1978; Domaax and Corey, 1978; Hakala, 1978; Morgan and Beeton, 1978; Grossniokle and Morgan, 1979; MordukhaiBoltovskoi, 1979.

Arctic Ocean Scott, 1899; Walton, 1922; Stephensen, 1933; Mob, 1953; Holmquist, 1958a, 1959a, 1973; McLaren, 1958; Grainger, 1962; Geiger, 1969; Just, 1970; Khlebovich et a,?.,1970; Zelickman and Golovkin, 1972; Sergeant, 1973; Prygunkova, 1974; Wiens et al., 1975; Ponomarenko, 1976, 1978; Wing and Ban, 1977; Hopkins and Gulliksen, 1978; Shih and Laubitz, 1978.

North Atlantic, north of 40"N, excluding the Baltic Sea Kinahan, 1859; Norman, 1865, 1880a, b ; Leslie and Herdman, 1881; Henderson, 1885; Walker, 1886, 1888, 189813; Scott, 1887, 1889, 1890, 1891a, b, 1894, 1896a, b, 1897, 1898, 1900a, b, 1902a, b ; Herdman, 1890, 1920; Herdman et al., 1898; Holt and Beaumont, 1899; Hjort andDahl, 1900; Meek, 1901,1903,1908,1909; Richard, 1904; Ostenfeld, 1906, 1916; Ostenfeld and Wesenberg-Lund, 1909; Mielk, 1911; Todd, 1914; McIntosh, 1919; Anonymous, 1920; Scott, 1922; Hardy, 1924; Pawsey and Davis, 1924; Sehodduyn, 1926; Broch, 1927; Percival, 1929; Savage, 1931; Otto and Wielinga, 1933; Stephensen, 1933, 1943; Moore, 1937; Sivertsen, 1938; Rees, 1939; Havinga, 1941; Redeke, 1941;Vos, 1941;Santo and Bacescu, 1942; Fraser and Saville, 1949a, b ; Fraser, 1950, 1956, 1961, 1970; Lochhead, 1950; Liao, 1951; Nouvel, 1951a, 1952, 1973c; Smidt, 1951; Jones, 1954; Le Sueur, 1954; Colman and Segrove, 1955; Fisher et al., 1955; Ostvedt, 1955; Anonymous, 1957; Bossanyi, 1957; Holmquist, 1957a, 1966, 1973; Labat, 1957; Brunel, 1959, 1960, 1961, 1963a, b, 1965, 1970; Raymont, 1961; Raymont and Conover, 1961; Rice, 1961; Mankowski, 1962; Molander, 1962; Prefontaine and Brunel, 1962; Salvat, 1962; Herman, 1963; Wigley, 1963; Raymont et al., 1964, 1966, 1967; Elofsson, 1965; Jepsen, 1965; Linford, 1965; Mauchline, 1965, 1967, 1968, 1969a, b, 1970a, b, 1971a, b, c, d, e, f, g, l972,1973a, b, c; Nagabhushanan, 1965; Almeida Prado, 1966; Jolicoeur and Brunel, 1966; MUUS, 1967; Richards and Riley, 1967; Edwards and Steele, 1968; Haefner, 1968, 1969; Da-hl and Mecklenburg, 1969; Geiger, 1969; O'Riordan, 1969; Miller, 1970; Schaefer, 1970; Baan and Holthuis, 1971; Barnes et a,?., 1971; Burton and Massie, 1971; McLusky and Heard, 1971; Wiborg, 1971; Brattegard and Vader, 1972; Kuhl and Mann, 1972; Lagardhre, 1972, 1977a, b ; Sage and Herman, 1972; Tyler, 1972, 1973; Morris,

366

THE BIOLOGY OF MYSIDS

1973; Morris and Sargent, 1973; Deniel, 1974; Hamond, 1974; Judkins and Wright, 1974; Williams'et al., 1974; Gmmatunga and Corey, 1975; Dadswell, 1975; Austin, 1976; McEachran et al., 1976; Moore, 1976; Moore and Moore, 1976; Nouvel and Lagardbre, 1976; Beaudouin, 1977; Boyden et al., 1977; Matthews and Bakke, 1977; Skjoldal and Bamstedt, 1977; Vlasblom and Elgershuizen, 1977; Warwick and Davies, 1977; Bamstedt, 1978; Foulds and Mann, 1978; Grabe, 1978; Hesthagen and Gjermundsen, 1978; MacDonald and Gilchrist, 1978; Makings, 1977, 1978; Schnack, 1978; Moore et al., 1979; Parker, 1979; Parker and West, 1979; Pezzack and Corey, 1979.

The Baltic Sea Suomalainen, 1954; SegerstrBle, 1957; Holmquist, 1959b; Lindquist, 1959; Wiktor, 1961; Mankowski, 1962; Shurh, 1962; Zmudzinski, 1962; Kuhlmorgen-Hille, 1963; Hmhtela, 1964, 1973; Siudzinski, 1964; Muus, 1967; Jansson and Kiillander, 1968; Uzars, 1969, 1972; Ackefors, 1969; Westin, 1970; Arntz, 1971, 1978a, b ; Zhuravel, 1972; Bagge and Ilus, 1973; Hesthagen, 1973; Nuorteva and Hiisiinen, 1975; Savolainen, 1975; Boysen, 1976; Hesthagen and Gjermundsen, 1979; Moller, 1979. The We-stern Atlantic, between 40"N and 40"s Whiteley, 1948; Fisher et al., 1955; Bowman, 1957, 1973, 1976, 1977b; Hulbert, 1957; Darnell, 1958; Hopkins, 1958, 1965, 1966; Gordon, 1960; Jander and Waterman, 1960; Bacescu, 1961, 1968b, C, d, 1969, 1970b; Evans, 1961; Steven, 1961; Cronin et al., 1962; Grant, 1962; Grice and Hart, 1962; Jander, 1962; Muncy, 1962; Price, 1962; Richards, 1963; Costa, 1964; Elofsson, 1965; Engel and Tan, 1966; Goodbody, 1965; Kohout and Kolipinsky, 1967; Emery, 1968; Herman et al., 1968; 0. S. Tattersall, 1968; Brattegard, 1969, 1970a, b, 1973, 1974a, b, 1975, 1977; Molenock, 1969; Markle and Grant, 1970; Silva, 1970a, b, 1972, 1974, 1977; Bacescu and Orghidan, 1971, 1977; Day et al., 1971; Wigley and Burns, 1971; Almeida Prado. 1972, 1973, 1974; Sikora et al., 1912; Swartz, 1972; Williams, 1972; Marsh, 1973; Quintero and Roa, 1973, 1977; Boesch and Dim, 1974; Haedrich and Henderson, 1974; Jacobs and Grant, 1974; Williams et al., 1974; Price, 1975, 1978; Stickney and Knowles, 1975; Stickney et al., 1975; Boesch et al., 1976; Holland and Polgar, 1976; McEachran et al., 1976; Maurer et al., 1976; Olge and Price, 1976; Camp et al., 1977; Moore and Sander, 1977; Ristich et al., 1977; Johnson and Hopkins, 1978; Overstreet and Heard, 1978; Price and Vodopich, 1979.

APPENDIX Ill

367

The Eaatern Atlantic, between 40"N and 40"S, but excluding the Mediterranean Sea and including South Africa Walker, 1888; Stebbing, 1905, 1908, 1914, 1917; Zimmer, 1912; Nouvel, 1941, 1973d; 0. S. Tattersall, 1957, 1958, 1961b, 1969; Belloc, 1959; Furnestin, 1959; Bainbridge, 1960; Bacescu and Mayer, 1961; Evans, 1961; Brown, 1964, 1976; Grindley and Penrith, 1965; Nouvel and Panouse, 1965; Day, 1967; Holthuis and Sivertsen, 1967; Bacescu, 1968a, 19730, 1976; Field, 1971; Brown and Talbot, 1972; Bacescu and Vasilescu, 1973; Merrett and Roe, 1974; Casanova, 1977; Christensen, 1978; Sargent et al., 1978; McLachlan et al., 1979.

The Mediterranean Sea Cerruti, 1921; Vannini, 1930; Bacescu, 1936a, 1966, 1970a; Caroli, 1937 ; Nouvel, 1951a, b, 1953, 1960; Delamare-Deboutteville, 1953a, b, c; Fisher et al., 1953; Hoenigman, 1954, 1958a, b, 1960, 1961, 1963, 1964a, b, 1965, 1968; Nouvel and Hoenigman, 1955; Genovese, 1956, 1957; Suau and Vives, 1957; Dion and Nouvel, 1960; Furnestin, 1960; Reys, 1960; Haidar, 1963; Ledoyer, 1963, 1965, 1966, 1974; Macquart-Moulin, 1965, 1970, 1973a, b, 1 9 7 7 ~ ;Vives, 1965, 1968; Bellan-Santini, 1966; Macquart-Moulin and Patriti, 1966; Casanova, 1968, 1970, 1977; Fishelson and Loya, 1968; Macquart-Moulin and Leveau, 1968; Champelbert and Macquart-Moulin, 1970; Franqueville, 1971; Moran and Fishelson, 1971; Moran, 1972; Morris, 1972; Ghirardelli, 1973; Bacescu and Schiecke, 1974; Hatzakis, 1974,1977; Kiortsis, 1974; Pesce, 1975a, b ; Wing, 1975; Ariani and Spagnuola, 1976; BacescuandMuradian, 1977; Cinelliet al., 1977; Wittmann, 1977,1978; Lucu, 1978.

The Black Sea Bacescu, 1938, 1954; Caspers, 1957; Valkanov, 1957; Zhuravel, 1959; Zatkutskiy, 1965, 1970; Dediu, 1966; Vinogradova and Petkevich, 1967; Nekrasova and Rakitina, 1968; Krylova et al., 1975; Savchuk, 1975.

The Caspian Sea Zenkevich, 1957 ; Vinogradova and Petkevich, 1967 ; Belyayeva et al., 1972; Zhuravel, 1972; Boroditch and Havlena, 1973; MordukhaiBoltovskoi, 1979.

The North Pacijic, north of 40"N Walker, 1898a; Shelford, 1935; Klawe, 1955; Tchindonova, 1959; &on, 1962; Pequegnat, 1965; Wing, 1965, 1975; Belyaev, 1966;

368

THE BIOLOUY OF MYSIDS

Hoffman and Yancey, 1966; Haertel and Osterberg, 1967; Miller, 1967; Jawed, 1969, 1973; Birstein' and Tchindonova, 1970; Green, 1970; Vinogradova, 1970; Vinogradova et al., 1970; Friedl, 1971a, b ; Pearcy and Ambler, 1974; Zelickman, 1974; Holmquist, 1975; Bordovskiy et al., 1976; Pearcy et al., 1977; Eggers et al., 1978.

The Eastern Pacijc between 40"N and 40"s Light, 1954; Holmquist, 195713, 1975; Clarke, 1961a; Heubach et al., 1963; Pequegnat, 1965; Ganssle, 1966; Painter, 1966; Radtke, 1966; Sasaki, 1966a, b, c; Stevens, 1966a, b ; Turner, 1966a, b ; Turner and Heubach, 1966; Bacescu, 1967, 1969, 1971a, b ; Clutter, 1967, 1969; Nakamura, 1967; Thomas, 1967; Childress, 1968, 1971a, b, 1975; Fager and Clutter, 1968; Adams, 1969; Heubach, 1969; Johnson, 1969; Hair, 1971; Simmons et al., 1974; Fuzessery and Childress, 1975; Kost and Knight, 1975; Maynard et al., 1975; Simmons and Knight, 1975; Belman and Childress, 1976; Hobson and Chess, 1976; Reel, 1978; Mickel and Chilress, 1978. The Western Pacijc between 40"N and 40"S, excluding Japanese coastal waters, China Seas and the Australasian region Russell, 1934; Dakin and Colefax, 1940; Hodge, 1963a, b, 1964; Nouvel, 1973b; Murano, 1974b, 1975, 1976, 1977a; Bell et al., 1978; Sale et al., 1978. Japanese coastal waters, Sea of Japan, China Sem and the Australasian region Ishikawa and Oshima, 1951; Matsudaira et al., 1952; Nouvel, 1957a, 1973a; 0. S. Tattersall, 1960a; Wickstead, 1961; Yamada, 1961; Fukai et al., 1962; Murano, 1963a, b, 1964a, b, 1966a, b, c, d, e, 1969, 1970a, b, c, 1974a, b, 1975, 1976, 1977a, b, 1978; Enomoto, 1964; Ii, 1964a, b ; -Ogawa, 1964; Kikuchi, 1967; Omori, 1969a; Taniguchi, 1969, 1976; Ikeda, 1970; Okada and Taniguchi, 1971; Shushkina,l972; Shushkina et al., 1972; Wing, 1975; Murano et al., 1976; Yamamoh, 1977; Sale et al., 1978. Eastern Indian Ocean, north of 40"s Alcock and Anderson, 1894; Nouvel, 1954a, b ; Pillai, 1961, 1963a, c, d, 1964, 1965, 1967, 1973; Wickstead, 1961; Balasubrahmanyan, 1964; Pillai and Mariamma, 1964; 0. S. Tattersall, 1965a; Kaliyamurthy, 1972; Shyamasundari, 1973; Gauld and Raju, 1975; Silas and Pillai, 1975; Balasubrahmanyan and Prince, 1976; Nath and Pillai, 1976; Taniguchi, 1976; Panampunnayd, 1977; Win, 1977.

APPENDIX Ill

369

Western Indian Ocean, north of 40"5 Alcock and Anderson, 1894; 0. S. Tattersall, 1958, 1962; Pillai, 1963b, 1965, 1968, 1973; Nouvel, 1964a, 1955, 1966b, 1967, 1969, 1971, 1974, 1978; Grindley and Penrith, 1965; Ledoyer, 1970; Ingle, 1972; Connell, 1974; Bacescu, 1975; Pratap et al., 1977; Christensen, 1978; McLachlan et al., 1979.

The Red Sea Nouvel, 1944; Bacescu, 1973a, b, d.

The Antarctic Ocean, south of 40"s Coutihre, 1917; Stephensen, 1927; Rustad, 1930; Mackintosh, 1934; Bary, 1956; Hodge, 1964; 0. S. Tattersall, 1961a, 1965; Birstein and Tchindonova, 1962; Ledoyer, 1969, 1977; Griffiths, 1976.

This Page Intentionally Left Blank

PART TWO THE BIOLOGY OF EUPHAUSIIDS

This Page Intentionally Left Blank

CHAPTER 1

INTRODUCTION The crustacean Order Euphausiacea contains 85 species distributed throughout the oceans of the world. A few species occur in coastal regions, a few in bathypelagic regions, but the majority live in the oceanic epipelagic and mesopelagic environments. They are easily recognized in plankton samples, many of the species being large in body size and classed as macroplankton or even as micronekton. The relatively small number of species in this Order has made it invaluable in biogeographical and oceanographical studies. I n addition, several species are becoming commercially important. A n assessment of the knowledge of the biology of euphausiids was published in 1969 as Volume 7 of this series (Mauchline and Fisher, 1969). It contains information that was available up to 1968. The present review assesses information published between 1968 and the end of 1978 but also includes some publications of 1979. An attempt has been made to present as complete a bibliography as possible, although the main purpose is t o assess, collate and synthesize the new information available. The present bibliography was closed during 1979 but many journals are issuing numbers behind schedule. Consequently, several relevant papers dated 1978 will have been omitted from the reference list. Mauchline and Fisher (1969), in their introduction, refer to a series of important papers that are not superseded by that work nor by the present one. They are: Sars (1885), Ruud (1932), John (1936), Einarsson (1945), Sheard (1953), Boden et al. (1955), Zimmer and Griiner (1956), Nemoto (1959, 1966), Brinton (1962), Marr (1962), Ponomareva (1963) and Mauchline (1967a). The systematics and distribution of species occurring in waters of south eastern Asia have been studied by Brinton (1975) who also presents revised maps of the world digtributions of several species. Commercial exploitation of the Antarctic Euphausia superba has stimulated Eddie (1977), Everson (1977) and Grantham (1977) to review the biology, distribution, harvesting and utilization of this species. Lomakina (1978) has written

374

THE BIOLOUY OF EUPHAUSIIDS

a systematic account of ,the Euphausiacea, in Russian, and collects together drawings of all the species in a single volume. The present review follows the format of Mauchline and Fisher (1969) as closely as possible t o facilitate cross-reference. An attempt is made to avoid duplication. Subjects are only discussed in the present work if they are new or alter conclusions presented in the earlier work. Additional material not affecting the earlier conclusions is presented as such, and usually briefly. The primary aims are t o produce, in conjunction with Mauchline and Fisher (1969), a source of data and information on this Order as well as t o identify areas where further investigation would be advantageous. The headings of the original subject index have been retained so that new information can be located quickly in the context of that presented in Volume 7. Omission of a subject heading indicates that no new information is available. There are, of course, many new topics that have been investigated in the last ten years and these are also listed. The taxonomy and synonymy of the species is as described by Mauchline and Fisher, with the exception of Thysanopoda aequalis/T. subaequalis where revision and description of a new species T . astylata have been effected. A number of minor nomenclatural changes have been made ; for example, Nyctiphunes couehii, Euphawia krohnii, Thysanoessa raschii and Stylocheiron suhmii have been modified to N . cowhi, E . krohni, T . raschi and S . suhmi respectively, to conform with modern usage.

CHAPTER 2

THE SPECIES OF KRILL In 1969 there were eighty-five valid species of euphausiids. There are still eighty-five because although a new species Thysanopoda astylata Brinton, 1975 has been described a former species, T . subaequnlis Boden, 1954 has been made a junior synonym of T . aequalis Hansen, 1905.

The external morphology of euphausiids is described in Mauchline and Fisher (1969) where there follows a historical account of the discovery and naming of the first known species. The Euphausiacea a t that time, was grouped together with the Mysidacea to form the Schizopoda. Next, the relationships of the euphausiids with other crustaceans were examined; Brinton (1966) remarks on the similarities between the Euphausiidae and the Sergestidae. The body form of a species can often vary, in some cases quite markedly. Mauchline and Fisher refer to descriptions of forms of Thysanoessa inermis, Nematoscelis dificitis, Stylocheiron afine and 8. longicorne. Reference is also made to smaller degrees of variation and to sexual dimorphism found among the following species : Euphausia diomedeae, E . triacantha, E . similis, E . vallentini, Nematoscelis microps, N . atlantica and N . tenella. New forms of species have been described and further accounts of variation within species are now available. Weigmann (1971) describes an isolated population of Pseudeuphusia latifrons in the Persian Gulf that is morphologically distinct from populations of this species in the Red and Arabian Seas. Adults in the latter regions are 6-12 mm (modal range, 7-9 mm) in body length while those in the Persian Gulf are 8-16 mm (modal range, 9-13mm). The petasma, the distal segment of the first pair of pleopods which is modified in males as a secondary sexual organ, is different in shape in males from the Persian Gulf when compared with males from the other two regions. The “neglecta” form of Thysanoessa inermis is described in some detail by Einarsson (1945) who concludes that it is predominantly a transitional form, occurring during the developiuental stages of the species, but can occasionally be retained by the mature adult. It can

376

THE BIOLOQY OF EUPHAUSIIDS

be recognized by examination of the second pair of thoracic legs but especially by the shape of the eye which is divided (bilobate) in the “neglecta” form and subcircular in the “ine~mis”form. Lacroix and Bourget (1973) have examined the forms of this species in the Gulf of St Lawrence and use an index of eye shape to quantify the tendency to have divided eyes. They measured the maximum width of the eye (LE2)and the width of the eye at the constriction (LEl). The index (EI) is

EI

LE2

=-

LE’

There is no significant difference in the shapes of the eyes between adult males and females of the same form. The eyes of the “neglecta” form are significantly more constricted than those of the “inermis” form. Further, they show that all the post-lmval and subadult individuals in that sea area have eyes of the “neglecta” form but that the “neglecta” form is more often retained in the mature adult stage than previously inferred by Einmsson; 8.1% of mature adults were of the “neglecta” form. Kulka and Corey (1978) and Berkes (1977b) confirmed this and the latter author also shows that there is a series of intermediate forms. The “neglecta” form of T . inermis does not appear to occur in the Pacific Ocean but seems to be restricted to the North Atlantic. Geiger et at, (1968) have found it in the Barents and Kara Seas. Thysanoessa inermis, as described by Einarsson (1945), has a small median dorsal spine at the posterior end of the sixth abdominal segment, the presence of which distinguishes this species from other North Atlantic species of the genus. Forms, however, with an additional spine on the fifth abdominal segment (two-spined form) and with additional spines on both the fourth and fifth segments (threespined form) were reported from the North Pacific and Bering Sea by Nemoto (1966) who reviews the earlier records. The one-spined form, usually the only form or dominant one present in a sea area, is present throughout the geographical range of the species. It is the dominant form in the Sea of Japan, minority form in the Bering Sea and Gulf of Alaska (Nemoto et al., 1973) the only form present in north-east Greenland, the Barents and Kara Seas (Geiger et al., 1968) and 6he dominant form in the fjords and coastal regions of western Norway (Wiborg, 1971). The two-spined form also appears to occur throughout most of the geographical range of the species. It is the minority form in the Sea of Japan, the dominant form off the eastern American coast (but a minority form in the Bay of Fundy

2. THE SPECIES OF KRILL

377

according to Kulka and Corey, 1978), North Sea and Pacific sector of the’hctic Ocean (Geiger et al., 1968) and a minority form in the fjords and coastal regions of western Norway (Wiborg, 1971). The three-spined form has not been reported since the description by Nemoto (1966). Zhuravlev (1977) describes the occurrence of spineless forms of Thysanoessa longpipes and Tessarabrachion oculatum in southern regions of the Sea of Okhotsk. There are two forms of Nematoscelis gracilis described by Gopalakrishnan (1975), the “old” form and the “new” form. The “old” form occurs in the northern part of the distributional area of the species, the “new” form in the southern part (Gopalakrishnan, 1974; Brinton, 1975). The “old” form is that of the type description; the “new” form differs from it in the morphology of the petasma, a significantly smaller adult body size, and the eyes in which the upper lobes are comparatively narrower. Gopalakrishnan (1975) plots the ratio of the length of the proximal process to the length of the median lobe of the petasma against carapace length and obtains a clear separation of these forms. The forms of Stylocheiron longiwrne and S . afline have been defined in the Pacific Ocean by Brinton (1962). There are two principal forms of 8.longicorne, a “long” form and a “short” form. A new N. Indian Ocean form, restricted to the northern Indian Ocean, is described by Brinton (1975). No information was previously available on the forms occurring in the Atlantic Ocean, except a single record of a male of the “short” form reported from the west coast of Scotland by Mauchline (1965). The forms are recognized biometrically through an examination of the proportions of the sixth abdominal segment in relation to the proportions of the eye (Brinton, 1962, 1975; Mauchline and Fisher, 1969). Baker (1970) found the “short” and “long” forms and intermediates between these present in the region of the Canary Islands. A similar investigation of populations off the coast of Portugal and in the Ligurian Sea produced the same result (Casanova, 1974). Consequently, there appears to be continuous variation within the Atlantic populabions and two distinct forms can not be separated. Baker (1970), however, found that the “long” form tends to occur at greater depths than the “short” form. All three forms occur in the Indian Ocean (Brinton, 1975). Baker (1970) examined S. afline from the region of the Canary Islands and found that they corresponded to three forms described by Brinton (1962) from the Pacific : West-Equatorial form, Central form and intermediates between these two. Brinton (1975) h d s that three forms occur in the Indian Ocean: the Indo-Australian form in

378

THE BIOLOGY OF EUPHAUSILDS

the north, the West-Equatorial form in central latitudes, and the Central form in southern latitudes. He suggests that the Central form occurs in the South Atlantic and that Baker’s samples are predominantly of the Central form, a conclusion not drawn by Baker nor supported by his data. There is sexual dimorphism in some species of the genus Nematoscelis, the males having an enlarged photophore or photophores and associated dorsal swellings in the abdominal segments (Mauchline and Fisher, 1969). Supplementary information is now available from Taniguchi (1967), James (1973) and Gopalakrishnan (1975) and is summarized in Table I. One form of N . microps has no enlarged photophores or dorsal swellings. The dorsal swellings of N . tenella and N . TABLEI. OCCURRENCE OF ENLARQED PHOTOPHORES (P)AND DORS- SWELLINQS (S) IN ABDOMINAL SEGMENTS OF MALESOF THE GENUSNemtoscelis

Abdominal eegmnt Species N . tenella N . microp8 N . atlantica N . gracilie

1

s

2

S

SP P

s

SP

3

4

S P

P

S P

P

P

atlantica are smaller and of a different form from those of the other two species (James, 1973 ; Gopalakrishnan 1975), although Taniguchi (1967) describes them as large in a Pacific specimen. James showed that the forms of each species tend to be separated geographically in bhe north-east Atlantic. James (quoted by Herring and Locket, 1978) found that, in the eastern Atlantic populations, male Nematobrachion jlexipes lack the third while females lack both the second and third abdominal photophores. The form of the petasmae, the modified first pair of pleopods of the male, is diagnostic of the species. It is difEcult to draw because it has to be unfolded, and therefore frequently distorted, but it is used successfully to identify the males. Identification of adult females of many species is still difficult and the potential usefulness of the female copulatory organs, the thelycum, was referred to in Mauchline and Fisher (1969). Recent work (Table 11) has confirmed its value. It is, however, even more difficult to illustrate and drawings of several

TABLE11. SPECIESIN Species Thysanopoda T . monacantha T . cristata T . tricuspidda T . aequalis T . astylata T . obtuaifrons T . pectinata T . orientalis T . microphthalrna T . acutifrons T . cornuta T . egregia T . spinicaudata Meganyctiphanes M . norvegica Nyctiphanes N . couchi N . australis N . capensis N . simplex Pseudeuphuaia P . latifrons P . sinica Eup h a d a E. americana E. eximia E. krohni E. mutica E. brevis E. diomedeae E. recurva E . superba E . vallentini E. lucens E. f rigida E. pacifica E. nana E. crystallorophias E. tenera E. similis E. similis var. armato E . mwronata E. sibogae E. distinguenda E. lamelligera E. gibba

WHICH THE

References 3, 8 8

3, 8 3, 5 8

8 1, 3, 8

1, 8 1, 2, 8 8 8

2, 5

5 5

3 4 4 4 4 4 4 4

3 9

THELYCWM HAS

BEEN

Species Euphada E. gibboides E. f a l h E. sanzoi E. pseudogibba E . paragibba E. hemigibba E. spinifera E . hanseni E . longirostris E . triacantha Tessabrachion T . oculatum Thysanoessa T . spinijera T . longipes T . inspinata T . inemnie T . longicaudatcc T . parva T . gregaria T . vicina T . nzacrura T . raschi Nenzatoscelis N . d i fl i l i s N . megalops N . tenellu N . microps N . atlantica N . lobata N . gracilis Nematobrmhion N . jlexipes N . sexspinosum N . boop's Stylocheiron S. carinaturn S. a$ne S. suhmi S . microphthalm S. insulare S. elongatum S. indicum S. Eongicorne S. abbreviatum S. ??aax,iTnum S. robuatum

DESCRIBED References

9 9 5, 9 4 4 4 4

7 7

2 2 7 6, 7 7 7 2

1 1, 5

3 1, 3

1, 5 1, 3

3 3

6

3, 6 6 6 6

References: 1, Einarsson, 1942; 2, Einarsson, 1946; 3, Sebastian, 1966; 4, Lomakine, 1972; 5, Costanzo and Guglielmo, 1976a; 6, Costanzo and Guglielmo, 1976b; 7, Costanzo and Guglielmo, 1977; 8, Guglielmo and Costanzo, 1977; 9, James, 1977.

380

THE BIOLOGY OF EUPRAUSIIDS

authors are compared in Fig. 1. Costanzo and Guglielmo (1976a b, 1977) use a scanning electron microscope and some distortion is present in some of the preparations illustrated in their photographs. This, however, is not a major problem. The greatest difficulty is in the production of instantly recognizable illustrations of this complex structure.

D Fxo. 1. The thelyca of: Thysanopodu orientaZw, A, after Eiriersson (19421, B, after Sebastian (1966); Meganytiphanes norvegica, C, after Costanzo and Guglielmo (1976a), D, after Einersson (1945);Stylocheimn longicorne, E, after Costanzo and Guglielmo (1970b), F, after Sebastian (1960).

A new taxonomic tool has been found by Mauchline and Nemoto (1977). There is a complex group of sensilla, present in the anterior

dorsal region of the carapace, usually located in the apical region of a keel or crest (Fig. 2). They illustrate the form of this organ in 48 species and demonstrate that the arrangement, proportions and shapes of the constituent elements are peculiar to each species. The organs are identical in males and females. The presence or absence of a lateral denticle on the carapace is sometimes used in the identification of species (Mauchline and Fisher,

2. THE

A

B

C

D

381

SPEOIES OF KRILL

E

F

G

H

FIQ.2. The position of the group of sensilla in the carapace of a euphausiid is indicated by the arrow. The patterns of the sensilla in ThysanoBsa species are shown, anterior end towards the top of the page: A, T. inermis; B, T . gregaria; C , T . raschi, N. E. Atlantic; D, T . raschi, N.W. Pacific; E, T . m c m r a ; F, T . longicaudata.; G , T . longipes and T . inspinata.; H, T . sphiifera (after Mauchline and Nemoto, 1977).

1969). McLaughlin (1965), referred to in Mauchline and Fisher (1969), described two forms of Nematoscelis dificilis, the typical form having no lateral denticle present; all the other characters of the two forms are identical. Recently, Talbot (1974) found an adult female Thysanopoda orientalis with a lateral denticle; it is normally only present in juveniles of this species and is absent in the adult. Variation in the occurrence of lateral denticles in Nematoscelis tenella is reported by James (1973). Thysanoksa raschi usually has one pair of lateral denticles present in the antexior region of the carapace. Mr Norman Nicol (personal communication) found two individuals in 6he North Sea (56'44'N lo47'E) with two pairs of lateral denticles, there being a smaller, but distinct, denticle a short distance posterior to the normal denticles. Re-examination of this species from Loch Fyne, Clyde Sea Area, confirmed the presence of such variant forms but they were not present in a sample of this species from British Columbia, Canada, kindly supplied by Dr J. Fulton. Hong (1969) compares the relative positions of the lateral denticles on the carapaces of Euphausia pacifica and E . n a m . Another character of taxonomic value, the pre-anal spine, is also known t o be variable. Mauchline and Fisher (1969) referred to Nemoto's discovery that the form of the pre-anal spine varied in Thysanohsa species. Hong (1969), Meira (1970) and Bartle (1976) refer to Nemoto's work and to variation in Thysanopoda tricuspidata, Nyctiphanes

382

THE BIOLOGY OF EUPHAUSIIDS

australis, Euphausia americana, E . eximia, E. pacijca, E . nana, E . hemigibba, Thysanoiksa longicaudata and Nematoscelis microps. The genetic variability of some euphausiids has been estimated by gel electrophoresis at 28-36 loci by Ayala et al. (1974, 1975) and Valentine and Ayala (1976). The Antarctic Euphausia superba had an observed frequency of heterozygous loci per individual of 5-8 & 0.4%, the temperate E . mucronata had 15.5+1.0% and the tropical E . distinguenda had 21.1 & 0.4%. Ayala et al. conclude that the genetic variability is low in regions where there is a pronounced seasonality of trophic resources while it is high in regions with stable resources. There is no correlation with the degree of stability of physical parameters of the environment. Lomakina (1968) compared Thysanopoda aequalis Hansen, 1905 with T . subaequalis Boden, 1954 and found intermediate forms. Baker (1 970) considered Lomakina’s evidence not substantial enough to cancel T . subaequulis. Brinton (1975) re-examined Hansen’s type material, relegated T . subaequalis to a junior synonym of T . aequalis, but created a new species T . astylata for individuals in which the third thoracic leg is not modified in either males or females, the dactylus being proportionate in size with the dactylus of the other thoracic legs. A comparison of the drawings of the thelyca of T . aequalis and T . subaequalis in Costanzo and Guglielmo (1976a, b) shows them to be virtually identical, thus reinforcing Brinton’s conclusions. There is confusion in the literature arising from the difficulties in separating Euphausia distinguenda afid E . sibogae (Sebastian, 1966; Brinton and Gopalakrishnan, 1973; Sawamoto, 1976) and this has probably not been fully resolved yet. Lomakina (1978) suggests that Pseudeuphausia sinica is a synonym of P . latifrons but this requires confirmation. Karedin (1 97 1 ) questions the species pair Nematoscelis dificilis and N . megalops but Gopalakrishnan (1975) suggests that they are both valid species. Mauchline and Nemoto (1977), having demonstrated the specific form of the compound organ in the carapace, found that those of Thysanohsa longipes and T . inspinata are identical; this result questions the status of the junior species, T . inspinata. Mauchline and Fisher (1969) pointed out the differences that seemed to exist between individuals of T . raschi from the north-east Atlantic and the Pacific; the compound organ in Atlantic animals is very different from that in individuals from the north-west Pacific (Fig. 2). The key to species of the genus Thysanopoda given in Mauchline and Fisher (1969) should be modified on p. 29 as follows: 6a. Frontal plate (rostrum) in lateral view tapers somewhat, distally.

2.

THE SPEUIES OF KRILL

383

Propodal and dactyl segments of endopodite of third thoracic leg of males normal .. .. .. .. .. .. astylata 6b. Frontal plate (rostrum) in lateral view, nearly of same thickness distally as proximally. Propodal segment of endopodite of third thoracic leg of males greatly reduced and dactylus modified as long naked spine .. .. .. .. .. .. aequalis Several supplementary keys are now available. A key to the species, adults and larvae, in the genus Nematoscelis is given by Gopalakrishnan (1975). Regional keys have been constructed: Indo-Pacific region by Brinton (1975), Gulf of Mexico by James (1970), Indian Ocean by Sebastian (1966), Arctic Basin and peripheral seas by Leung (1970), and the north Atlantic (Mauchline, 1971a). Lomakina (1978) provides a key t o the Euphausiacea in Russian. Pertzova (1976) provides a key to larvae of the Antarctic species, Casanova-Soulier (1968) to the furciliae and Casanova (1972) to the calyptopes of Mediterranean species.

CHAPTER 3

DISTRIBUTION AND SYNONYMY Much more information is now available regarding the distribution of species in the Indian Ocean and waters of south-east Asia. This has resulted primarily from the International Indian Ocean Expedition 1959-1965 but also from other investigations such as the Naga expedition 1959-1961 (Brinton, 1975). The majority of the geographical distributions of the species, illustrated on maps by Mauchline and Fisher (1969), have not been altered substantially. The presence of a species has frequently been confirmed in sea areas where few records were previously available. Some new investigations of the distributions of species in the Atlantic Ocean have been published but more information is still required, especially in the central and southern Atlantic. Brinton (1975) has revised the world distributions of more than 40 species and some of 6hese maps have been redrawn here because they are significantly different from those given by Mauchline and Fisher (1969). The format of the following list follows that of Mauchline and Fisher (1969). The name of the species is followed by sources of any new descriptions of the adults. Extensions to or major qualifications of the geographical distribution of the species are then presented. This is followed by an additional list of references to the distribution of the species published since 1968. A few important references prior to 1968 were omitted by Mauchline and Fisher ( 1969) and these have now been included. One new species, Thysanopoda astylata Brinton, 1975, has been added to the list. Family BENTHEUPHAUSIIDAE Genus Bentheuphausia G. 0. Sars, 1885 Bentheuphausia amblyops G. 0. Sam, 1885 No new descriptions. Present throughout the Gulf of Mexico (James, 1970). (Roger, 1968c, d, 1974b, c, 1978a; O’Riordan, 1969; Ponomareva, 1975, 1976c.)

3.

DISTRIBUTION AND SYNONYMY

385

Family EUPHAUSIIDAE Genus Thysanopoda Milne-Edwards, 1830 Thysanopoda monacantha Ortmann, 1893 Brinton, 1975: 162-164, figs 87, 119g.

New records of this species in the Atlantic and Indian Oceans are available and Brinton (1975) has produced a new map of its distribution. James (1970) has found it throughout the Gulf of Mexico and it occurs in the Atlantic between about 40"N and 2O-3O0S, as shown in Mauchline and Fisher (1969). Present in the Indian Ocean between l0"N and 20"s (Brinton, 1975) but does not occur in the Arabian Sea and Bay of Bengal as shown by Ponomareva (1975). (Sheard, 1965; Sebastian, 1966; Brinton, 1967a, 1979; Roger, 1968c, d, 1974b, c, 1978a, Gopalakrishnan and Brinton, 1969; Karedin, 1969b;Weigmann, 1970; Brinton and Gopalakrishnan, 1973; De Decker, 1973; Talbot, 1974; Legand et al., 1975; Youngbluth, 1975; Ponomareva, 1976c; Taniguchi, 1976; Weigmann-Haass, 1976; McWilliam, 1977; Hu; . 1978; Rottman, 1978.)

Thysanopoda cristata Q. 0. Sars, 1883 Brinton, 1975: 160, figs 86, 1190.

Brinton (1975) has produced a map of its distribution that includes the new records from the Atlantic (Baker, 1970) and the Gulf of Mexico (James, 1970). Widespread in the Atlantic Ocean between 35"N and 30'5, in the Indian Ocean between 10°N and 30'5. Distribution in the Pacific Ocean is similar to that shown in Mauchline and Fisher (1969). (Roger, 1968c, d, 1974b, c, 1978a; Gopalakrishnan and Brinton, 1969; Brinton and Gopalakrishnan, 1973; Legand et al., 1975; Youngbluth, 1975; Ponomareva, 1976c; McWilliam, 1977; Rottman, 1978.)

Thysanopoda tricwpidata Milne-Edwards, 1837 Brinton, 1975; 155-156, figs 81, 119a.

Brinton (1975), in a revised map, indicates its occurrence in the north-west Atlantic to about 45"N and includes the records of James (1970) from the Gulf of Mexico. Distribution in Indian and Pacific Oceans as shown in Mauchline and Fisher, 1969. (Sebastian, 1966; Roger, 1968c, d, 1974b, c, 1978s; Gopalakrishnan and Brinton, 1969; Karedin, 1969b; Meira, 1970; Weigmann, 1970, 1974b; Brinton and Gopalakrishnan, 1973; De Decker, 1973; Talbot, 1974; Legand et al., 1975; Ponomaseva, 1975, 1976c; Youngbluth, 1975; Andreu, 1976;

386

THE BIOLOGY OF ECJPHAUSIIDS

Taniguchi, 1976; McWilliam, 1977; Moore and Sander, 1977; Rottman, 1978.)

Thysanopoda aequalis Hansen, 1905 T . aequalis Hansen, 1905: 18-20; 1910: 84-85, pl. xii, fig. 4a-b ( ? c), pl. xiii, fig. l a ; Zimmer, 1914: 417-418, figs 53, 54; Tattersall, 1924: 15-16 pl. ii, figs 5, 6; Brinton, 1975: 165-166, figs 89a, b, d, e, 119, k, 1; T . subaequalis Boden, 1954: 190-192, fig. 5; Boden, Johnson and Brinton, 1955: 297; Boden and Brinton, 1957: 337-341, figs 1-2; Bacescu and Mayer, 1961: 180-190, fig. 4a; James, 1970: 226; Baker, 1970: 311-313; Weigmann, 1974b: 23-24

Thysanopoda subaequalis Boden, 1954 is now considered a junior synonym of T . aeqwlis Hansen, 1905 (see Brinton, 1975). Brinton states that T . aequalis occurs in the Atlantic Ocean, Mediterranean Sea, Indian Ocean south of the equator and in the central and western Pacific Ocean between 40"N and the equator as shown in Fig. 3. Thysanopoda astylata Brinton, 1975 T.astylata Brinton, 1975: 168-169, figs 80 d-f, 119n, 0. T. aequalie Boden, 1954: 190, fig. 4c-f; Boden, Johnson and Brinton, 1955: 303-305, fig. 9; Boden and Brinton, 1957: 337-340, figs la, 2c; Brinton and Gopalakrishnan, 1973: 368-369, fig. 9a, b.

This species is absent from the Atlantic Ocean and Mediterranean Sea. It is present in the Indian Ocean north of about 5"s but absent from the Arabian Sea and northern region of the Bay of Bengal. It occurs in the eastern Pacific between 40"N and 10°N and across the southern Pacific from about 10"N to 35'5, as shown in Fig. 3. (Brinton, 1967a, 1973, 1975; Brinton and Wyllie, 1976.) NOTE: Brinton considers references to the distribution of T . aequalis and T . subaequalis in the following papers to be of T. aequalis: T . aequalis: Casanova, 1974; T . subaequalis: Brinton and Gopalah4shnan, 1973; Casanova, 1974. Brinton believes the references t o the distribution of T . aequalis or T . subaequalis in the following papers refer to either that of T . aequalis or T . astylata: Weigmann, 1970; Roger, 1973a. The following papers present information on the distribution of T . aequalis and T . subaequalis but are not considered by Brinton: Macquart-Moulin and Leveau, 1968; T . aequalis Sebastian, 1966; Brinton, 1967a, 1973; Roger, 1968c, d, 1974b, c, 1978a; Gopalakrishnan and Brinton, 1969; Casanova, 1970; Casanova-Soulier, 1970a; Weigmann, 1970; Franqueville, 1971; Owre and Foyo, 1972; Wiebe and D'Abramo, 1972; Casanova et al., 1973; Talbot, 1974; Legand et al., 1975; Ponomareva, 1975, 1976c; Youngbluth, 1975; Taniguchi, 1976; McWilliam, 1977; Moore and Sander, 1977; Lindley, 1977;

1200

160°

160°

1200

800

40°

O0

40°

~

FIG.3. Distributionof Thysanopoda a e q d k and T.astylata (after Brinton, 1975).

80°

388

THE BIOLOGY OF EUPHAUSIIDS

Sipos, 1977; Hu, 1978; Rottman, 1978. T . subaequalis Roger, 1968c, 1974c; Gopalakrishnan and Brinton, 1969; Casanova, 1970; Talbot, 1974; Youngbluth, 1975; Andreu, 1976.

Thysanopoda obtusifrons G. 0 . Sam, 1883 Brinton, 1975: 169-170, fig. 90, 119m.

A revised map of the distribution of this species is provided by Brinton (1975) and is essentially the same as that in Mauchline and Fisher (1969). Atlantic Ocean: occurs between 40"N and 30'5, including the Gulf of Mexico but excluding (no records so far) the regions of the Guinea and south-eastern Atlantic basins. Indian Ocean: Brinton (1975) shows that it occurs between the equator and 30"S, while the distribution shown by Ponomareva (1976) is incorrect. Pacific Ocean : as previously shown by Mauchline and Fisher (1969). (Brinton, 1967a; Roger, 1968c, d, 1974b, c, 1978a; Gopalakrishnan and Brinton, 1969; Karedin, 1969b; Baker, 1970; James, 1970; Weigmann, 1970, 1974b; Brinton and Gopalakrishnan, 1973; De Decker, 1973; Legand et al., 1975; Youngbluth, 1975; Ponomareva, 1976c; Taniguchi, 1976; McWilliam, 1977.)

Thysanopoda pectinata Ortmann, 1893 Brinton, 1975: 162, figs 86, 119f.

Brinton (1975), in a revised map of its distribution, shows it occurring widely in the Atlantic Ocean between 46"N and 30'5, in the Indian Ocean between l0"N and 40"s. Its distribution in tihe Pacific is as shown in Mauchline and Fisher (1969). (Sebastian, 1966; Brinton, 1967a, 1979; Roger, 1968c, d, 1974b, c, 1978a; Gopalakrishnan and Brinton, 1969; Baker, 1970; James, 1970; Brinton and Gopalakrishnan, 1973; De Decker, 1973; Legand et al., 1975; Youngbluth, 1976; Ponomareva, 1976c; Taniguchi, 1976; Hu, 1978; Rottman, 1978.)

Thysanopoda orientalis Hansen, 1910 Brinton, 1975: 164, figs 88, 119h.

Distribution in the Pacific Ocean as shown in Mauchline and Fisher (1969). Occurs in Indian Ocean between the equator and 40"s and in the south Atlantic Ocean between about 20"s and 40"s ; it also occurs in the Gulf of Mexico and off the eastern coast of the United States (Brinton, 1975) but has not been recorded from the central and eastern Atlantic between latitudes 40"N and lO0-20"S. (Sebastian, 1966; Roger, 1968c, d, 1974b, c, 197th; Gopalakrishnan and Brinton, 1969; James, 1970; Brinton and Gopalakrishnan, 1973; De Decker,

3.

DISTRIBUTION A?XD SYNONYMY

389

1973; Talbot, 1974; Ponom,areva, 1975, 1976c; Taniguchi, 1976; McWilliam, 1977; Rottman, 1978; Brinton, 1979.) Thysanopoda microphthalma G. 0 . Sms, 1885 Brinton (1975) in a revised map, shows this species restricted to the Atlantic Ocean south of 40"N but absent from the south-western Atlantic. The records shown in Mauchline and Fisher (1969) for this species in the Indian Ocean are not commented on by Brinton. No record of this species from %heAgulhas Current is given by Talbot (1974). (Baker, 1970.) Thysanopoda acutifrons Holt and Tattersall, 1905 A few questionable records north of 20"s in the Indian Ocean are given by Mauchline and Fisher (1969); it is illustrated as a transitional zone species, occurring between 20"s and 40"s. Ponomareva (1975), however, shows it present throughout the Indian Ocean between about 18"N and 20"s but not in the region 20"s t o 40"s. It is not mentioned by Brinton and Gopalakrishnan (1973) as present in this region of the Indian Ocean. (Wiborg, 1968; Jones, 1969; O'Riordan, 1969; Kawamura, 1974a; Talbot, 1974; Jorgensen and Matthews, 1975; Bmtle, 1976; Ponomareva, 1976c; Lindley, 1977; Robertson and Roberts, 1978.) Thysanopoda cornuta Illig, 1905 Brinton, 1975: 166-158, figs 82, 119b.

There is still not enough information on this bathypelegic species to define its distribution. (Roger, 1968c, 1974c, 197%; Brinton and Gopalakrishnan, 1973; Brinton, 1975.) Thysanopda egregia Hansen, 1905 Brinton, 1975: 158-160, figs 84, 119d.

James (1970) has recorded it from the north-eastern region of the Gulf of Mexico. (Roger, 1968c, 1974c, 1978a; Brinton and Gopalakrishnan, 1973; Brinton, 1975; Youngbluth, 1975; Ponomareva, 1976c; Rottiman, 1978.) Thysanopoda spinicaudata Brinton, 1953 Brinton, 1975: 168, figs 83, 119c.

Recorded at 13'36" Gopalakrishnan, 1973.)

65'03'E in the Arabian Sea. (Brinton and

390

THE BIOLOGY OF EUPHAUSIIDS

Genus Meganyctiphanes Holt and Tattersall, 1905 Meganyctiphanes noivegica (M. Sars, 1857) The distribution of this species is shown in Mauchline and Fisher (1969). Its occurrence in the Barents Sea is restricted mainly to the western areas. (Glover, 1952; Southward, 1962; Lie, 1967; Geiger et al., 1968; Macquart-Moulin and Leveau, 1968; Vives, 1968; Guglielmo, 1969; Jones, 1969; O'Riordan, 1969; Baan and Holthuis, 1970; Casanova, 1970, 1974; Casanova-Soulier, 1970a; Fraser, 1970; Franqueville, 1971; Wiborg, 1971; Arbault et al., 1972; Bainbridge and Forsyth, 1972; Bouchet and Thiriot, 1972; Lagardere, 1972, 19778, b ; Wiebe and D'Abramo, 1972; Matthews, 1973; Mauchline, 1973; Sergeant, 1973; Beaudouin, 1974; Hollingshead and Corey, 1974; Soboleva, 1974; Weigmann, 1974b; Berkes, 1976; Degtyaryova et al., 1976; Kashkin, 1976; Le ROUX,1976; Boyden et al., 1977; Elder and Fowler, 1977; J.-P. Casanova, 1977; Johannessen, 1977; Lindley, 1977, 1978b; Matthews and Bakke, 1977; Sipos, 1977; Bliznichenko et al., 1978; Hopkins et al., 1978.) Genus Nyctiphanes, G. 0. Sars, 1883 Nyctiphanes couchi (Bell, 1853) The distribution in the north-eastern Atlantic (Mauchline and Fisher, 1969) extends from tihe northern coast of mainland Scotland north-westiwards to 60°N l0"W and north-eastwards to the Shetland Islands; it does not occur as far north as the Faroes. It is also present sporadically off the northern and western coasts of Ireland. There is no evidence of breeding populations in the Irish Sea (Khan and Williamson, 1970) although the species occurs in this area. (Glover, 1952; Southward, 1962; Baan and Holthuis, 1969; O'Riordan, 1969; Casanova-Soulier, 1970a; Casanova, 1970, 1974; Fraser, 1970; Lagardere, 1972, 1977a, b ; Hesthagen, 1973; Mauchline, 1973; Beaudouin, 1974, 1977; Jorgensen and Matthews, 1975; Le ROUX,1976; Lindley, 1977, 1978b; Sipos, 1977; Gros and Cochard, 1978.)

Nyctiphanes australis G. 0 . Sars, 1883 Its distribution is as shown in Mauchline and Fisher (1969). (Sheard, 1965, 1967; Knox, 1970; Jillett, 1971; Bradford, 1972; Jillett and Mitchell, 1973; Bartle, 1976; Jillett, 1976; Markina, 1976.) Nyctiphanes capensis Hansen, 1911 Mauchline and Fisher (1969) show this species restricted to South African coastal regions. This has now been shown to be incorrect. Meira (1970) found it present at the Cape Verde Islands and Andreu

3.

DISTRIBUTION AND SYNONYMY

391

(1976) and Weigmann-Haass (1976) record it present in the region of Cape Blanc, north-west Africa. It was not present off the Ivory Coast (Binet, 1976). (Brinton and Gopalakrishnan, 1973; De Decker, 1973; Talbot, 1974, Cram and Schulein, 1974.)

Nyctiphanes simplex Hansen, 1911 The distribution of this species is shown in Mauchline and Fisher (1969). (Brinton, 1967a, 1973, 1979; Antezana, 1970; O'Connell 1971 ; Brinton and Wyllie, 1976; Kanaeva and Pavlov, 1976.) Genus Pseudeuphausia Hansen, 1910 Pseudeuphausia latifrons (G. 0. Sam, 1883) Brinton, 1975: 172, figs 91, 121h.

Brinton (1975) has produced a revised map of the distribution of this species but does not include the information by Ponomareva (1975). It is present off eastern Madagascar northwards to the Arabian Sea and Gulf of Oman (including the Red Sea) then south-eastwards to the equator in the region south of India, through the Bay of Bengal and from there south-eastwards to north-western Australia. The other distributional areas are shown in Mauchline and Fisher (1969). (Sebastian, 1966; Sheard, 1967 ; Gopalakrishnan and Brinton, 1969 ; Hong, 1969; Weigmann, 1970, 1971 ; Brinton and Gopalakrishnan, 1973; Taniguchi, 1976; McWilliam, 1977; Rottman, 1978.)

Pseudeuphausia sinica Wang and Chen, 1963 No additional information is available t o that given in Mauchline and Fisher (1969). Lomakina (1978) suggests that this species is a junior synonym of P . latifrons. Genus Euphausia Dana, 1852 Euphausia americana Hansen, 19 1 1 The area of distribution is as shown in Mauchline and Fisher (1969) except that it has now been recorded in the south-west Atlantic near 32"s 50"W (Ramirez, 1971, 1973, 1977). (Kelly and Dragovich, 1968; Baker, 1970; James, 1970; Meira, 1970; Owre and Foyo, 1972; Weigmann-Haass, 1976; Moore and Sander, 1977.)

Euphausia eximia Hansen, 19 1 1 Brinton and Wyllie, 1976: fig. 13.

The distribution remains as shown in Mauchline and Fisher (1969). Bacescu (1975) states that he found this species occurring with the mysids Pseudanchialina erythraea and a species of Gastrosaccus in the

392

THE BIOLOGY OF EUPEAUSIIDS

sub-littoral region of the coast of Tanzania, East Africa, at a depth of 1-6 m ; this record requ’ires confirmation. (Brinton, 1967a, 1973, 1979; Kinzer, 1969b; Meira, 1970; Roger, 1974c, 1978a.)

Euphausia krohni (Brandt, 1851) Mayer, 1967: 16-20, figs 1-3.

The distribution of this species is as shown in Mauchline and Fisher (1969) except that it has now been recorded as far north as 62-63”N, off the western coast of Norway. (Glover, 1952; Macquart-Moulin and Leveau, 1968; Razouls and Thiriot, 1968; Vives, 1968; Wiborg, 1968; Guglielmo, 1969; Jones, 1969; O’Riordan, 1969; Baker, 1970; Casanova, 1970, 1974; Casanova-Soulier, 1970a; Fraser, 1970; Franqueville, 1971; Bouchet and Thiriot, 1972; Lagardere, 1972, 1977a, b ; Wiebe and D’Abramo, 1972; Beaudouin, 1973, 1974; Kiortsis, 1974; Andreu, 1976; Weigmann-Haass, 1976; Lindley, 1977; Sipos, 1977.)

Euphausia rnutica Hansen, 1905 Brinton, 1975: 176-177, figs 92, 120e.

Brinton (1975) has produced a revised map of its distribution, essentially the same as in Mauchline and Fisher (1969), with the addition of records of its presence in the Gulf of Mexico (James, 1970). (Sheard, 1965; Brinton, 1967a, 1973; Roger, 1968c, 1974b, c, 1978a; Gopalakrishnan and Brinton, 1969; Hong, 1969; Karedin 1969b; Kinzer, 1969b; Weigmann, 1970, 1974b; Brinton and Gopalakrishnan, 1973; De Decker, 1973; Talbot, 1974; Ponomareva, 1975; Youngbluth, 1975; Andreu, 1976; Taniguchi, 1976; Moore and Sander, 1977; Lindley, 1977; MeWilliam, 1977; Rottman, 1978.)

Euphausia brevis Hansen, 1905 Brinton, 1975: 177-178, figs 93, 120b.

The revised map of its distribution, drawn by Brinton (1975), is essentially the same as in Mauchline and Fisher (1969) except that James (1970) has found this species present in the Gulf of Mexico. Ponomareva (1975)wrongly shows it occurring throughout the Arabian Sea and Bay of Bengal. (Mayer, 1967; Sebastian, 1966; Roger, 1968c, d, 197413, c , 1978a; Gopalakrishnan and Brinton, 1969; Karedin, 1969b; Baker, 1970; Casanova, 1970, 1974; Casanova-Soulier, 1970a; Weigmann, 1970, 1974b; O w e and Foyo, 1972; Wiebe and D’Abramo, 1972; Brinton and Gopalakrishnan, 1973; Casanova et al., 1973; De Decker, 1973; Talbot, 1974; Youngbluth, 1975; Andreu, 1976; Taniguchi, 1976; Lindley, 1977; MeWilliam, 1977; Reid, 1977.)

3. DISTRIBUTION AND SYNONYMY

393

Euphausia diomedeae Ortmann, 1894 Brinton, 1975: 180, figs 95, 120d.

The revised map of ,Brinton (1975) is almost the same as that in Mauchline and Fisher (1969) except in two regions. Brinton shows an extension westwards round the southern regions of South Africa and another extension of the distribution northwards in the western Pacific to off western Japan. It does not occur south of 20"s in the eastern Indian Ocean as shown by Ponornareva (1975). (Sebastian, 1966; Brinton, 1967a, 1979; Gopalakrishnan and Brinton, 1969; Weigmann, 1970, 1974a; Wiebe, 1972; Brinton and Gopalakrishnan, 1973; Casanova et al., 1973; De Decker, 1973; Kawamura, 1973, 1974a; Roger, 1974b, c, 1978a; Talbot, 1974; Ponomareva, 1976c; Taniguchi, 1976; McWilliam, 1977; Reid, 1977; Rottman, 1978.)

Ewphawia recurva Hansen, 1905 Brinton, 1975: 178, figs 94, 12013.

The revised map of its distribution given by Brinton is identical with that in Mauchline and Pisher (1969) except that Illig's record, off north-west Africa, now considered incorrect, has been omitted. The distribution in the South Pacific is not so well d e h e d , as indicated by Brinton (1975), and Bartle (1976) has recently summarized records of its occurrence h the Tasman Sea-New Zealand region. (Sheard, 1966, 1967; Brhton, 1967a, 1973, 1979; Karedin, 1969b; Antezana, 1970; Ramirez, 1971, 1973, 1977; Brinton and Gopalakrishnan, 1973; De Decker, 1973; Kawamura, 1973; Talbot, 1974; Markina, 1976; Taniguchi, 1976; McWilliam, 1977; Nepgen, 1977; Rottman, 1978.)

Euphausia superba Dana, 1850 The distribution of this species is as shown in Mauchline and Fisher (1969). Antezana et al. (1976a) have discovered a population in a fjord in southern Chile at approximately 50"s. (Beklemishev, 1960; Sasaki et al., 1968; Bogdanov et al., 1969; Shevtsov and Makarov, 1969; Makarov and Shevtsov, 1969, 1971; Bogdanov and Solyanik, 1970; Ivanov, 1970; Makarov et at., 1970; Mackintosh, 1972, 1973; Makarov, 1972, 1974b, 1976; Moroz, 1973; Kawamura, 1974a; Voronina, 1974; Korabel'nikov, 1975; Latogurskii et al., 1975; Makarov and Maslennikov, 1975; Antezana et al., 197633; Everson, 1976, 1977; Freytag, 1977; Mohr and Fischer, 1977; Hempel and Hempel, 1978; Kock and Stein, 1978; Pommeranz, 1978; Sarhage et al., 1978.)

394

THE BIOLOGY OF E U P H A U S D S

Euphausia vallentini Stebbing, 1900 Distribution is as shown in Mauchline and Fisher (1969). It has also been recorded, by Bradford (1972) a8 a southern visitor to waters of New Zealand. (Lomakina, 1966; Antezana, 1970; Knox, 1970; Ramirez, 1971, 1973, 1977; Kawamura, 1970, 1974a; Reid, 1977; Robertson and Roberts, 1978.) Euphuusia lucens Hansen, 1905 Distribution is as described in Mauchline and Fisher (1969). (Sheard, 1965; Lomakina, 1966; Antezana, 1970; Ramirez, 1971, 1973, 1977; Bradford, 1972; De Decker, 1973; Kawamura, 1974; Talbot, 1974; Bartle, 1976; Nepgen, 1977; Reid, 1977; Robertson and Roberts, 1978.) Euphausia frigida Hansen, 1911 Information is available, additional to that in Mauchline and Fisher (1969), especially of the region of the Weddell Sea and West Wind Drift. (Kawamura, 1974; Makarov, 1977; Hempel and Hempel, 1978.) Euphausia pacijica Hansen, 191 1 Brinton and Wyllie, 1976: fig. 15.

Its distribution is shown in Mauchline and Fisher (1969). It is recorded incorrectly from the coast of Burma in the Indian Ocean by Win (1977). (Brinton, 1967a, 1973; Hong, 1969; Karedin, 1969b; Taniguchi, 1969; Day, 1971; Fried, 1971a, b ; O’Connell, 1971; Kawamura, 1973; Fedotova, 1976; Gardner, 1977; Reid, 1977; Shvetsova, 1977; Zhuravlev, 1977.)

Euphausia nana Brinton, 1962 Its distribution is shown in Mauchline and Fisher (1969). (Hong, 1969.)

Euphausia crystallorophias Holt and Tattersall, 1906 Its distribution is shown in Mauchline and Fisher (1969) and there is no additional information available. Euphausia tenera Hansen, 1905 Brinton, 1975: 190, figs 100, 120f.

Brinton (1975) provides a revised map of the distribution of this species. The map is essentially the same as that in Mauchline and Fisher (1969) but includes its occurrence in the Gulf of Mexico reported by James (1970). This species does not occur in the Arabian Sea and

3. DISTRIBUTION

AND SYNONYMY

395

Bay of Bengal as shown by Ponomareva (1975). (Sheard, 1965;Sebastian, 1966; Brinton, 1967a, 1979; Roger, 1968c, 1974b,c, 1978a; Gopalakrishnan and Brinton, 1969; Hong, 1969; Karedin, 1969b; Kinzer, 1969b;'Lewisand Fish, 1969; Weigmann, 1970; Owre and Foyo, 1972; Wiebe, 1972; Brinton and Gopalakrishnan, 1973; De Decker, 1973; Kawamura, 1973; Talbot, 1974; Ponomareva, 1975; Youngbluth, 1975; Andreu, 1976; Taniguchi, 1976; WeigmannHaass, 1976;Moore and Sander, 1976, 1977;Lindley, 1977;McWilliam, 1977; Reid, 1977; Rottman, 1978.)

Euphausia similis G. 0. Sam, 1883 Brinton, 1975: 180-181, figs 96, 120e.

The revised map of its distribution in Brinton (1975) is closely similar to that in Mauchline and Fisher (1969). Ponomareva (1975), however, shows the main concentration of this species to be present in the western central Indian Ocean, but this requires confirmation. (Sheard, 1965, 1967 ; Lomakina, 1966; Gopalakrishnan and Brinton, 1969; Hong, 1969; Karedin, 1969b; Weigmann, 1970; Ramirez, 1971, 1973, 1977; Bradford, 1972; De Decker, 1973; Brinton and Gopalakrishnan, 1973;Bartle, 1976;Taniguchi, 1976;Robertson and Roberts, 1978; Rottman, 1978.)

Euphuusia similis var. armata Hansen, 1911 The distribution is as shown in Mauchline and Fisher (1969). (Sheard, 1965, 1967; Lomakina, 1966; Antezana, 1970; De Decker, 1973; Kawamura, 1974; Brinton, 1975; Bartle, 1976; Markina, 1976; Robertson and Roberts, 1978.) Euphausia mucronata G. 0. Sars, 1883 The distribution is as shown in Mauchline and Fisher (1969). (Antezana, 1970; Reid, 1977; Brinton, 1979.) Euphausia sibogae Hansen, 1908 Brinton, 1975: 188-190, figs 99, 120g; Sawamoto, 1976: 66-67, fig. 3a,c.

Mauchline and Fisher (1969) show this species restricted to the East Indian Archipelago and off north-east Australia. Brinton (1975), following Sebastian (1966), considers that the population in the northern Indian Ocean of the closely related species E . distinguenda is in fact not of this species but of a form more closely related to E . sibogae. He has consequently applied the name E . sibogae to this population of E . distinguenda and re-defines its distribution as in Fig. 4. Sawamoto (1976)has found E . sibogae in the sea area to the

1200

160'

160°

1200

80°

40°

40°

BOO

60'

40'

20' 00 20c

4OC

6OC

I I

1200

160°

160°

1 I

1200

800 FIG.4. Distribution of Epcphazlsia sibogae and 1.&ti-

I I

40°

00

40°

(after Brinton, 1975).

I

800

3.

DISTRIBUTION AND SYNONYMY

397

north of New Guinea, off Taiwan and off southern Japan (Fig. 4). Ponomareva (1975) shows 'its distribution in the Indian Ocean incorrectly. (Brinton and Gopalakrishnan, 1973, as "Euphausia distinguenda" Group. As E . distinguenda : Reuben, 1968; Gopalakrishnan and Brinton, 1969; Weigmann, 1970; Casanova et al., 1973; Taniguchi, 1976; &William, 1977.)

Euphausia distinguenda Hansen, 1911 Sawamoto, 1976: 68-69, fig. 3b, d.

According to Brinton (1975) this species is restricted to the tropical region of the Eastern Pacific (Fig. 4). The populations shown occurring in the Indian Ocean in Mauchline and Fisher (1969) have been renamed E. sibogae (see above under E . sibogae). (Roger, 1978a; Brinton, 1979.)

Euphausia lamelligera Hansen, 1911 Its distribution is as shown in Mauchline and Fisher (1969). (Brinton, 1967a, 1979; Wiebe, 1972.) Euphausia gibba G. 0. Sars, 1883 Brinton, 1975: 185, figs 98e, 121m.

Mauchline and Fisher (1969) and Brinton (1975) show this species with a distribution restricted to the South Pacific between about 15"s and 30"s. Sawamoto (1976), however, has found two males and two females at 0°18-9'N, 133"357'E, the area north of New Guinea. (Sheard, 1965, 1967; Roger, 1974b, 1978a; Youngbluth, 1975.)

Euphausia gibboides Ortman, 1893 Brinton and Wyllie, 1976: fig. 14.

Brinton (1975) has produced a revised map of the distribution of this species (Fig. 5). Its distribution in the Atlantic Ocean is shown as very different from that indicated by Mauchline and Fisher. It is also shown as not occurring in the Indian Ocean, previous records being ascribed to the closely related species E . sanzoi. (Brinton, 1967a, 1973, 1979; Karedin, 1969b; Antezana, 1970; Baker, 1970; James, 1970; Pugh, 1972; Roger, 1974c, 1978a; Weigmann, 1974b; Andreu, 1976; Weigmann-Haass, 1976.)

Euphausia fallax Hansen, 1916 Brinton, 1975: 184-185, figs 97g-i, 121f, g.

Brinton (1975) has revised the distribution of this species along with that of the closely related E. sanzoi. It occurs in the Indo-Australian region and off north-east Australia; it has also been recorded

60°--

--60(

I

i

1200

160°

i

160° FIQ.6. Distribution of Euph-

i i

i

500

fi

i

i

40° pWmkka, E . fal2az and E . a a d (after Brinton, 1975).

1200

40°

00

i

800

3. DISTRIBUTION

AND SYNONYMY

399

from the Andaman Islands, off southern Ceylon, and off northern Madagascar (Fig. 5). More information is required to define its distribution. (Roger, 1968c, 1974b, c, 1978a; Brinton and Gopalakrishnan, 1973;Knight, 1978.)

Euphausia sanzoi Torelli, Brinton, 1975: 182-184, figs 97+f, 121, c-e.

1934

Brinton (1975) has re-defined the distribution of this species. This species has been confused with the two closely similar species E . gibboides and E . fallax. Brinton now thinks that it is E . sanzoi that occurs in the Red Sea and northern Arabian Sea and extends southwards along the western margin of the Indian Ocean to Madagascar (Fig. 5). It also occurs off south-western Japan, through the SouCh China Sea and sporadically to north-eastern Australia and the region bounded by latitudes 1O0S-30"S off Australia and longitudes 170°W-1500E (Fig. 5). (Gopalakrishnan and Brinton, 1969; Weigmann, 1970, 1974a; Brinton and Gopalakrishnan, 1973; Rottman, 1978.)

Euphausia pseudogibba Ortmann, 1893 Brinton, 1975: 186, figs 98a,b, 121h,i.

Brinton (1975) has revised the distribution of this species (Fig. 6). He shows it as confined t o the tropical Atlantic, with an extension in the Gulf Stream, present acrosa the northern Indian Ocean but not in the Arabian Sea or northern region of the Bay of Bengal, and present in the Indo-Australian region. The distributional map in Ponomareva (1975) is wrong, as is the description of its distribution in Ponomareva (1972). Its distribution in the Pacific remains as shown in Mauchline and Fisher (1969). (Sebastian, 1966; Gopalakrishnan and Brinton, 1969; James, 1970; Weigmann, 1970; Brinton and Gopalakrishnan, 1973; Andreu, 1976; Taniguchi, 1976; WeigmannHaass, 1976; McWilliam, 1977; Moore and Sander, 1977; Rottman, 1978; Brinton, 1979.)

Euphausia paragibba Hansen, 1910 Brinton, 1975: 186, figs 98d, 121k,1.

Brinton (1975) shows the.distribution of this species to be the same as that illustrated in Mauchline and Fisher (1969). Ponomareva (1975) shows this species widely distributed throughout the Indian Ocean but this is incorrect. (Roger, 1968c, 1974c; Gopalakrishnan and Brinton, 1969; Weigmann, 1970; Brinton and Gopalakrishnan, 1973; De Decker, 1973; Talbot, 1974; Ponomareva, 1976c; Taniguchi, 1976; Brinton, 1979.)

v,

n I

I

I 20°

I 60°

I I

160°

I I

1200

I

800

I

40°

FIG.6. Distribution of Euphawia pseudogibba (after Brinton, 1975).

3.

DISTRIBUTION AND SYNONYMY

401

Euphausia hemigibba Hansen, 1910 Brinton, 1976: figs QSc, 121j.

The revised distribution .*hewn in Brinton (1975) is essentially the same as in Mauchline and Fisher (1969). Ponomarevrt (1976) incorrectly extends its distribution in the eastern’ Indian Ocean to the south-eastern areas of the Bay of Bengal. (Brinton, 1967a, 1973, 1979; Mayer, 1967; Macquart-Moulin and Leveau, 1968; Vives, 1968; Guglielmo, 1969; Karedin, 1969b; Baker, 1970; Casanova, 1970, 1974; Casanova-Soulier, 1970a; James, 1970; Meira, 1970; Owre and Foyo, 1972; Wiebe and D’Abramo, 1972; Brinton and Gopalakrishnan, 1973; Casanova et al., 1973; De Decker, 1973; Talbot, 1974; Weigmann, 1974b ; Andreu, 1976; Taniguchi, 1976; Weigmann-Haass, 1976; Lindley, 1977; McWilliam, 1977; Sipos, 1977.)

Euphausia spinifera G. 0 . Sam, 1883 Distribution is as shown in Mauchline and Fisher (1969). (Sheard, 1965,1967;Lomakina, 1966; DeDecker, 1973; Kawamura, 1974; Talbot, 1974; Barhle, 1976; McWilliam, 1977; Robertson and Roberts, 1978.)

Euphausia hanseni Zimmer, 1915 Its distribution is shown in Mauchline and Fisher (1969). (De Decker, 1973; Andreu, 1976; Weigmann-Haass, 1976, 1977.)

Euphausia longirostris Hansen, 1908 Its distribution is shown in Mauchline and Fisher (1969). (Reid, 1977; Robertson and Roberts, 1978.)

Euphausia trimantha Holt and Tattersall, 1906 Its distribution is shown in Mauchline and Fisher (1969). (Hempel and Hempel, 1978.) Genus Tessarabrachion Hansen, 1911 Tessarabrachion oculatum Hansen, 1911 Its distribution is shown in Mauchline and Fisher (1969). (Karedin, 1969b; Taniguchi, 1969; Day, 1971; Brinton, 1973; Zhuravlev, 1977.) Genus Thysano&sa Brandt, 1851 Thysanoessa spinifera Holmes, 1900 Brinton and Wyllie, 1976: fig. 20.

Its distribution is shown in Mauchline and Fisher (1969). (Brinton, 1967a, 1973; Day, 1971; Fried, 1971a, b ; Kawamura, 1973; Peterson and Miller, 1975; Hobson and Chess, 1976.)

402

THE BIOLOGY O F EUPHAUSIIDS

Thysano&sa longipes Brandt, 1851 Its distribution is as shown in Mauchline and Fisher (1969) except that it has now been recorded from the eastern region of the Beaufort Sea and north of the Bering Strait. (Geiger et al., 1968; Hong, 1969; Karedin, 1969b; Taniguchi, 1969; Day, 1971; Brinton, 1973; Fedotova, 1976; Zhuravlev, 1976b, 1977; Shvetsova, 1977; Wing and Barr, 1977.)

ThysanoQsa inspinata Nemoto, 1963 Its distribution is as shown in Mauchline and Fisher (1969). (Karedin, 1969b; Taniguchi, 1969.) Thysanoika inermis (Krrayer, 1846) The distribution of this species is as shown in Mauchline and Fisher (1969). (Glover, 1952; Lie, 1967; Geiger et al., 1968; Bedard, 1969; Jones, 1969; O’Riordan, 1969; Fraser, 1970; Wiborg, 1971; Bainbridge and Forsyth, 1972; Zelickman and Golovkin, 1972; Kawamura, 1973; Lacroix and Bourget, 1973; Nemoto et al., 1973; Sergeant, 1973; Soboleva, 1974, 1975; Jorgensen and Matthews, 1975; Berkes, 1976; Degtyaryova et al., 1976; Drobysheva and Soboleva, 1976; Fedotova, 1976; Zhuravlev, 197613, 1977; Drobysheva et al., 1977; Lindley, 1977, 1978b; Matthews and Bakke, 1977; Reid, 1977; Shvetsova, 1977; Wing and Barr, 1977; Bliznichenko et al., 1978; Hesthagen and Gjermundsen, 1978; Hopkins et al., 1978; Kulka and Corey, 1978; Schnack, 1978; Shih and Laubitz, 1978.)

Thysanoessa longicaudata (Krrayer, 1846) Its distribution is shown in Mauchline and Fisher (1969) but Meira (1970) has now recorded it as far south as the Cape Verde Islands. (Glover, 1952; Geiger et al., 1968; Jones, 1969; O’Riordan, 1969; Zelickman and Golovkin, 1972; Beaudouin, 1073, 1974, 1977; Soboleva, 1974, 1975; Jorgensen and Matthews, 1975; Berkes, 1976; Degtyaryova et al., 1976; Drobysheva et al., 1977; Lindley, 1977, 1978a, b ; Matthews and Bakke, 1977; Bliznichenko et al., 1978; Sameoto, 1978.)

Thysanoiissaparva Hansen, 1905 Its distribution is shown in Mauchline and Fisher (1969). (Baker, 1970; De Decker, 1973; Kawamura, 1974.) Thysanoessa gregaria G. 0. Sars, 1883 Brinton and Wyllie, 1976: fig. 19.

Its distribution is shown in Mauchline and Fisher (1969). (Sheard, 1965, 1967; Lomakina, 1966; Brinton, 1967a, 1973, 1979; Jones,

3.

DISTRIBUTION AXD SYNONYMY

403

1969; Karedin, 1969b; O'Riordan, 1969; Antezana, 1970; Baker, 1970; Casanova, 1970, '1974; Casanova-Soulier, 1970a; Knox, 1970; Rmirez, 1971, 1973, 1977; Bradford, 1972; Brinton and Gopalakrishnan, 1973; De Decker, 1973; Kawamura, 1974; Talbot, 1974; Weigmann, 1974b; Andreu, 1976; Bartle, 1976; Jillett, 1976; WeigmannHaass, 1976; Lindley, 1977; McWilliam, 1977; Reid, 1977.)

Thysanoessa vicina Hansen, 1911 The distribution of this species is shown in Mauchline and Fisher (1969). Ramirez (1971) found it in the south-west Atlantic near 50'5, 55"W.(Kawamura, 1974.) Thysanoessa macrura G. 0. Sam, 1883 Its distribution is shown in Mauchline and Fisher (1963). (Kawamura, 1974.) Thysano&sa raschi (M. Sam, 1864) Its distribution is shown in Mauchline and Fisher (1969) but it has been found by Geiger et al. (1968) in the Arctic Basin at almost 80"N. (Glover, 1952; Bedard, 1969; Jones, 1969; Micklukhina, 1969; Fraser, 1970; Day, 1971; Wiborg ,1971; Zelickman and Golovkin, 1972; Hesthagen, 1973; Mauchline, 1973; Sergeant, 1973; Prygunkova, 1974; Soboleva, 1974, 1975; Wing, 1974; Jorgensen and Matthews, 1975; Berkes, 1976; Degtyaryova et al., 1976; Drobysheva and Soboleva, 1976; Fedotova, 1976; Zhuravlev, 197613, 1977; Drobysheva et al., 1977; Lindley, 1977, 197813; Shvetsova, 1977; Wing and Barr, 1977; Bliznichenko et al., 1978; Hopkins and Gulliksen, 1978; Hopkins et al., 1978; Shih and Laubitz, 1978.) Genus Nematoscelis G. 0. Sam, 1883 Nematoscelis dificilis Hansen, 1911 Brinton and Wyllie, 1976: fig. 16.

Its distribution is shown in Mauchline and Fisher (1969). (Brinton, 1967a, 1973, 1979; Karedin, 1969b; O'Connell, 1971; Day, 1971; Kawamura, 1973; Gopalakrishnan, 1974.)

Nematoscelis megalops G. 0. Sam, 1883 The distribution is shown in Mauchline and Fisher (1969) and Gopalakrishnan (1974). It has been recorded in the Atlantic as far north as the North Cape at 71"N and also at 70"N 14"E and 75"N 19"E. (Glover, 1952; Sheard, 1965; Lomakina, 1966; Maver, 1967; MacquartMoulin and Leveau, 1968; Razouls and Thiriot, 1968; Vives, 1968;

404

THE BIOLOGY OF EUPHAUSIIDS

Wiborg, 1968; Guglielmo, 1969; Jones, 1969; O'Riordan, 1969; Antezana, 1970; Baker, 1970; Casanova, 1970, 1974; Casanova-Soulier, 1970a; Fraser, 1970; Knox, 1970; Franqueville, 1971; Ramirez, 1971, 1973, 1977; Bouchet and Thiriot, 1972; Bradord, 1972; Owre and Foyo, 1972; Weibe and D'Abramo, 1972; Brinton and Gopalakrishnan, 1973; De Decker, 1973; Kawamura, 1974; Talbot, 1974; Weigmann, 1974b; Jorgensen and Matthews, 1975; Andreu, 1976; Bartle, 1976; Weigmann-Haass, 1976; Lagardere, 1977a, b ; Lindley, 1977; Sipos, 1977; Boyd et ul., 1978; Robertson and Roberts, 1978; Wiebe and Boyd, 1978.)

Nematoscelis tenella G. 0 . Sam, 1883 Brinton, 1975: 192-194, figs 101, 122a.

The distribution of this species shown by Brinton (1975)is different from that of Mauchline and Fisher (1969) only in the South Atlantic. Brinton shows it present throughout the region north of 40"N whereas Gopalakrishnan (1974) shows its distribution to be similar t o that shown by Mauchline and Fisher but has added seven new records in the south-eastern Atlantic. He suggests that it may be present in the south-west Atlantic but it was not found there by Ramirez (1971, 1973, 1977). It does not occur in the northern regions of the Arabian Sea and Bay of Bengal as illustrated by Ponomareva (1975). (Sebastian, 1966; Brinton, 1967a, 1973, 1979; Roger, 196% d, 1974b, C, 1978a; Gopalakrishnan and Brinton, 1969; Karedin, 1969b; Baker, 1970; James, 1970; Weigmann, 1970, 1974b; Brinton and Gopalakrishnan, 1973; De Decker, 1973; James, 1973; Talbot, 1974; Ponomareva, 1976c; Youngbluth, 1975; Taniguchi, 1976; Weigmann-Haass, 1976; McWilliam, 1977; Rot%man,1978.)

Nematoscelis microps G. 0. Sam, 1883 Brinton, 1975: 195, figs 103, 122c.

The distribution as mapped in Mauchline and Fisher (1969), Gopalakrishnan (1974) and Brinton (1975) is closely similar. Ponomareva (1975) shows this species present throughout the Arabian Sea and Bay of Bengal whereas other authors show it absent from these regions with the exception of the eastern half of the Bay of Bengal. (Sheard, 1965, 1967; Sebastian, 1966; Mayer, 1967; Roger, 1968c, d, 1974b, C, 1978a ; Gopalakrishnan and Brinton, 1969; Karedin, 1969b; Baker, 1970; James, 1970; Meira, 1970; Brinton and Gopalakrishnan, 1973; De Decker, 1973; James, 1973; Talbot, 1974; Weigmann, 197413; Youngbluth, 1975; Ponomareva, 1976c; Taniguchi, 1976; WeigmannHaass, 1976; McWilliam, 1977; Rottman, 1978; Brinton, 1979.)

3.

DISTRIBUTION AND SYNONYMY

405

Nemutoscelis atlantica Hansen, 1910 Brinton, 1975: 195-196, figs 104, 122d.

Gopalakrishnan (1974) defines the distribution of this species in the Indian Ocean where it had previously been unknown (Mauchline and Fisher, 1969) (Fig. 7). I t occurs in the central water mass in latitudes 15’5 to 40”s. Ponomareva (1975) incorrectly maps this species as occurring north of the equator in the Indian Ocean and even being present in the Arabian Sea. Gopalakrishnan (1974) records it, for the &st time, present in the North Atlantic gyre at about 50”N. (Brinton, 1967a, 1973, 1975; Razouls and Thiriot, 1968; Roger, 1968c,d 1974b, c, 1978a; Gopalakrishnan and Brinton, 1969; Karedin, 1969b; Baker, 1970; Casanova, 1970, 1974; Casanova-Soulier, 1970a; James, 1970; Bouchet and Thiriot, 1972; Wiebe and D’Abramo, 1972; Brinton and Gopalakrishnan, 1973; De Decker, 1973; James, 1973; Weigmann, 1974b; Youngbluth, 1975; Ponomareva, 1976c; McWilliam, 1977; Sipos, 1977; Rottman, 1978.)

Nematoscelis lobata Hansen, 1916 Brinton, 1975: 196-198, figs 104c, d.

Gopalakrishnan (1974) added seven new records to the two in Mauchline and Fisher (1969). All records of this species are restricted t o the seas around the western side of the Philippines (Fig. 7). (Karedin, 1969b; Brinton, 1975.)

Nematoscelis gracilis Hansen, 1910 Brinton, 1975; 194, figs 102, 122b.

The distributional map in Mauchline and Fisher (1969) has now been altered t o take account of the two forms of this species known to exist (Fig. 8). The distributional area of the species remains unaltered but one form, known as the “old” form, inhabits the northern region of the distribution while the obher form, the “new” form, inhabits the southern regions (Gopalakrishnan, 1974; Brinton, 1975). (Sebastian, 1966; Brinton, 1967a, 1979; Roger, 1968c, 1974b, c, 1978a; Gopalakrishnan and Brinton, 1969; Karedin, 1969b; Antezana, 1970; Brinton and Gopalakrishnan, 1973; De Decker, 1973; Kawamura, 1973; Ponomareva, 1975, 1976c; Youngbluth, 1975; Taniguchi, 1976; Reid, 1977; McWilliam, 1977; Rottman, 1978.) Genus Nematobrachion Calman, 1905 Nematobrachion jlexipes (Ortmann, 1893) Brinton, 1975: 198-200, figs 105, 102f.

The distribution of this species mapped by Brinton (1975) differs

~ L U .I .

usmbution of N ~ ~ ~ ~ t oatkzntica s ~ e l i sand N . lobata (after Brinton, 1975).

t

6ooL n I

1

I

1:00

160°

1

1

I

1200

80°

I

40°

1

O0

I

40’

FIG.8. Distribution of the “Old” and “New” forms of Nematoacelis gracilis (after Brinton, 1975).

I

80’

408

THE BIOLOQY OF EUPHAUSIIDS

from that in Mauchline and Fisher (1969) in the following regions. The species is now known to extend into the South China Sea. It does not occur in the Gulf of Oman (Brinton, 1975; Ponomareva, 1975). It is widespread throughout the Indian Ocean between about 10"N and 40'5 and not as shown by Ponomareva (1975). There are additional records in the Atlantic off northwest Africa and in the Gulf of Mexico. (Sebastian, 1966; Brinton, 1967a, 1973, 1979; Roger, 1968c, d, 1974b, c, 1978a; Gopalakrishnan and Brinton, 1969; Antezana, 1970; Baker, 1970; James, 1970; Weigmann, 1970, 1974b; Day, 1971; Brinton and Gopalakrishnan, 1973; De Decker, 1973; Talbot, 1974; Youngbluth, 1975; Andreu, 1976; Ponomareva, 1976c; Taniguchi, 1976; Weigmann-Haass, 1976; McWilliam, 1977.)

Nernatobrachion sexspinosum Hansen, 1911 Brinton, 1975: 200-202, figs 106, 122g.

Brinton (1975) has produced a revised map of the distribution of this species (Fig. 9). There is no change in the limits of its distribution in the Pacific Ocean, described by Mauchline and Fisher (1969), but much more knowledge is now available about its distribution in the Atlantic and, especially, Indian Oceans. Brinton (1975)shows it occurring in two regions of the Atlantic, one in the north between latitudes 35"N and 20"N, the other in the south between about 20'5 and 30"s. Baker (1970) and Weigmann (1974b) both record it off north-west Africa. Indian Ocean: it occurs between the equator and about 30's in the central Indian Ocean (Fig. 9) and does not have the distribution shown by Ponomareva (1975). (Roger, 1968c, d, 197413, c, 1978a; Karedin, 1969b; James, 1970; Brinton and Gopalakrishnan, 1973; Youngbluth, 1975; Ponomareva, 1976c; Taniguchi, 1976; McWilliam, 1977; Hu, 1978.)

Nematobrachion boopis (Calman, 1905) Brinton, 1975: 202, figs 107, 122h.

Brinton (1975) maps the distribution of this widely ranging species but there is litble change from the map given in Mauchline and Fisher (1969). It occurs to about 50'5 in the Indian Ocean and does not have the much more restricted distribution suggested by Ponomareva (1975). (Sebastian, 1966; Roger, 1968c, d, 1974b, c, 1978a; O'Riordan, 1969; Baker, 1970; Fraser, 1970; James, 1970; Brinton and Gopalakrishnan, 1973; Ponomareva, 1976c; Youngbluth, 1975; Andreu, 1976; Taniguchi, 1976; Weigmann-Haass, 1976.)

-60 I

120°

I

160°

1

I

160°

1

I

120°

I

800

FIG.9. Distribution of Nemaf~bm.&hnexspi-

I I

40°

I

O0 (after Brinton, 1976).

40° 1

800 1

410

THE BIOLOGY OF EUPEAUSIIDS

Genus Stylocheiron G. 0.Sars, 1883 Stylocheiron carinaturn, G. 0 . Sam, 1883 Brinton, 1976: 204-206, figs 108, 123a.

The revised distribution of this species (Brinton, 1975) is virtually unchanged from that shown in Mauchline and Fisher (1969). (Sheard, 1965, 1967; Sebastian, 1966; Brinton, 1967a, 1973, 1979; Roger, 1968c, d, 1974b, c, 197th; Karedin, 1969b; Gopalakrishnan and Brinton, 1969; Baker, 1970; James, 1970; Weigmann, 1970, 1974b; Owre and Foyo, 1972; Wiebe, 1972; Brinton and Gopalakrishnan, 1973; Casanova et al., 1973; De Decker, 1973; Talbot, 1974; Ponomareva, 1975, 1976c; Youngbluth, 1975; Taniguchi, 1976; WeigmannHaass, 1976; McWilliam, 1977; Moore and Sander, 1977; Rottman, 1978.)

Stylocheiron afine Hansen, 1910 Brinton, 1976: 208-210, figs 111, 123d-f; Brinton and Wyllie, 1976: fig. 18.

Brinton (1975) has re-defined the distributions of the various forms of this species in the Pacific and Indian Oceans (Figs 10 and 11). The distributions in the Pacific remain the same as those shown in Mauchline and Fisher (1969), except that the West-Equatorial form has now been shown to extend northwards in the western Pacific in the Kuroshio Current. Only the western-Equatorial form is shown to occur in the Indian Ocean in Mauchline and Fisher (1969). Brinton (1975) shows this form to occur in a band across the central Indian Ocean between about l0"N and 18"-20"S. The Indo-Australian form, as well as occurring extensively in the Indo-Australian region with a northern extension of this distribution in the Kuroshio, also occupies the northern Indian Ocean, from the equator northwards through the Arabian Sea t o the Gulfs of Aden and Oman and in the greater part of the Bay of Bengal. The Central form occurs on the southern side of the distributionak area of the West Equatorial form, being distributed across the Indian Ocean between latitudes 15"-17"S and 30"s. Brinton does not provide any further information on the distribution of this species in the Atlantic Ocean. Ramirez (1971) has now found it in the south-west Atlantic near 31"s 48"W. (Sebastian, 1966; Brinton, 1967a, 1973; 1979; Roger, 1968c, 197413, c, 19788; Gopalakrishnan and Brinton, 1969; Hong, 1969; Karedin, 196913; Antezana, 1970; Baker, 1970; James, 1970; Weigmann, 1970, 1974a, b ; Wiebe, 1972; Brinton and Gopalakrishnan, 1973; Casanova et al., 1973; De Decker, 1973; Talbot, 1974; Ponomareva, 1975; Youngbluth, 1975; Taniguchi, 1976; McWilliam, 1977 ; Ramirez, 1977 ; Rottman, 1978.)

1200

160°

160°

rrm

40'

1200

40°

80"

form

50°

t60C

n

I

I

I

1

1

1

120°

160'

160°

1200

I

1

8 0 '

I

40°

I

00

I 40°

I 80°

FIG.10. Distribution of the West-Equatorialform, East-Equatorialform and California Current form ofStylocheirm aflne (afterBrinton, 1975).

--60

...........:..... ...

I "

U

"

O

v,

Central form

n

L

I I

1200

I

160°

1

1

160°

I

I

1200

I

800

I I

400

I

0

I

40°

800

Fra: 11. Distribution of the Indo-Australianform 8nd Centre1 Form of Stylochiron aflne (after Brinton, 1975).

3.

DISTRIBUTION AND SYNONYMY

413

Stylocheiron suhmi G. 0. Sam, 1883 Brinton, 1975: 206, figs 109, 123b.

Brinton (1975) presents a revised map of the distribution of this species that differs little from that in Mauchline and Fisher (1969). There are small additional regions of occurrence shown in the equatorial Pacific but the species is less widely distributed in the IndoAustralian region than indicated in Mauchline and Fisher. Brinton shows it restricted between the equator and about 30"s in the Indian Ocean, whereas Ponomareva (1975) incorrectly shows it present throughout the Arabian Sea and southern half of the Bay of Bengal. It is present in the Atlantic Ocean between latitudes 40"N and 30'5. (Sheard, 1965; Brinton, 1967a, 1973; Mayer, 1967; Vives, 1968; Gopalakrishnan and Brinton, 1969; Karedin, 1969b; Kinzer, 1969b; Casanova, 1970, 1974; Casanova-Soulier, 1970a; James, 1970; Weigmann, 1970, 1974b; Owre and Foyo, 1972; Wiebe and D'Abramo, 1972; Brinton and Gopalakrishnan, 1973; Casanova et al., 1973; De Decker, 1973; Roger, 1974b, 1978a; Talbot, 1974; Youngbluth, 1975; Taniguchi, 1976; Weigmann-Haass, 1976; McWilliam, 1977; Pratap et al., 1977; Sipos, 1977; Rottman, 1978.)

Stylocheiron microphthulma Hansen, 1910 Brinton, 1975: 210-212, figs 112, 123g.

According t o Brinton (1975) it is not so widely distributed in the northern Indo-Australian region as indicated in Mauchline and Fisher (1969) but it only occurs in the southern half of this region. It extends from there westwards across the Indian Ocean between latitudes 5"N to 10-20"s (Brinton, 1975; Ponomareva, 1975). There we no new records of its occurrence, in addition to the two quoted in Mauchline and Fisher (1969), in the Atlantic Ocean, and it is probable that these records were erroneous and the species does not occur in the Atlantic. (Sebastian, 1966; Gopalakrishnan and Brinton, 1969; Karedin, 1969b; Weigmann, 1970; Brinton and Gopalakrishnan, 1973; De Decker, 1973; Roger, 1974b, 1978a; Talbot, 1974; Taniguchi, 1976; McWilliam, 1977; Rottman, 1978.)

Stylocheiron insulare Hansen, 1910 Brinton, 1975: figs 110, 123c.

Brinton has mapped the distribution of the species (Fig. 12). It occurs in the Indo-Australian region, extending eastwards in the equatorial Pacific t o about 170"E. It has also been found in the Indian Ocean in the region of the Andaman Islands and off southern Sumatra

60°

I I 1200

I

I

160°

I 160°

1

I

1200

I 80°

I

I

40°

I

I

I

O0

40'

80°

FIG.12. Distribution of Stylocheiron ineu&re and S. indicum (after Brinton, 1976).

3.

DISTRZBlJTION AND SYNONYMY

415

and Java. There are no recent records of it occurring in the Atlantic Ocean. (Brinton and Gopalakrishnan, 1973.)

Stylocheiron elongaturn G. 0. Sam, 1883 Brinton, 1975; 214, figs 114, 123.

The revised map of the diskibution given by Brinton (1975) is closely similar t o that in Mauchline and Fisher (1969) except that Brinton does not show it occurring in the Gulf of Oman. It does not occur in the Gulfs of Aden and Oman and the western Arabian Sea, as suggested by Ponomareva (1975). (Sebastian, 1966; Roger, 1968c, d, 1974b, c, 1978a; Baker, 1970; Fraser, 1970; James, 1970; Beaudouin, 1973, 1974; Gopalakrishnan and Brinton, 1973; De Decker, 1973; Talbot, 1974; Weigmann, 1974b; Youngbluth, 1975; Bartle, 1976; Taniguchi, 1976; Weigmann-Haass, 1976; McWilliam, 1977; Reid, 1977; Brinton, 1979).

Stybcheiron indicum Silas and Mathew, 1967 Brinton, 1976: 212-214, figs 113, 123h.

Brinton (1975) shows bhe three regions where this species has been found (Fig. 12): the type locality off the west coast of India, the western Philippines and the Moluccas. The distribution of this species given by Brinton and Gopalakrishnan (1973) is wrong, according to Brinton (1975). (Gopalakrishnan and Brinton, 1969.)

Stylocheiron longicorne G. 0 . Sam, 1883 Brinton, 1975: 216-216, figs 116, 124a-c.

Brinton (1975) maps the distribution of three forms of this species (Fig. 13): the “long” form, the “short” form and the North Indian Ocean form. The distributions of the “long” and “short” forms in the Pacific Ocean remain unchanged from those in Mauchline and Fisher (1969) except that both forms have been shown to occur in the South China Sea. Mauchline and Fisher plotted the distribution of the species, form unspecified, in the Atlantic and Indian Ocean. Brinton (1975) shows one form, the North Indian Ocean form, present north of about 5”N in the western Arabian Sea and Gulf of Oman and also in the western Bay of Bengal. The “short” form occurs throughout the Indian Ocean between latitudes 5”N and 30”s and connects via the southern half of the Indo-Australian region with Pacific populations. The “long” form occurs between latitudes 5”N and 40”s in the Indian Ocean and is also connected through the same region with Pacific populations. The distribution of the “short” form is not continuous westwards round South Africa although it may be

Erratum Figures 13 and 14 on pages 416 and 418 are incorrectly positioned and should be transposed.

3. DISTRIBUTION AND SYNONYMY

417

present in the central south Atlantic. The "long" form is present mound South Africa and thoughout the Atlantic Ocean as far north as 60"N. Ramirez (1971, 1977) has found this species, form not defined, in the south-west Atlantic near 31'5, 48"W, while James (1970) records the species throughout the Gulf of Mexico. (Sebastian, 1966; Brinton, 19678, 1973, 1979; Macquart-Moulin and Leveau, 1968; Roger, 1968c, d, 1974b, c, 1978a; Vives, 1968; Gopalakrishnan and Brinton, 1969; Karedin, 1969b; O'Riordan, 1969; Antezana, 1970; Baker, 1970; Casanova, 1970, 1974; Casanova-Soulier, 1970a; F'raser, 1970; James, 1970; Weigmann, 1970, 197413; Franqueville, 1971; Owre and Foyo, 1972; Wiebe and D'Abramo, 1972; Brinton and Gopalakrishnan, 1973; De Decker, 1973; Talbot, 1974; Youngbluth, 1975; Ponomareva, 1976c; Taniguchi, 1976; Andreu, 1976; WeigmannHaass, 1976; Beaudouin, 1977; Lindley, 1977; McWilliam, 1977; Moore and Sander, 1977; Sipos, 1977; Rottman, 1978.)

Stylocheiron abbreviatum G. 0 . Sam, 1883 Brinton, 1976: 220, figs 118, 124f.

The revised distribution map of Brinton (1975) is almost identical with that of Mauchline and Fisher (1969), except that the species is now known to occur in the Red Sea and Gulf of Eilat and also throughout the Gulf of Mexico. It does not occur in the northern Arabian Sea as suggested by Ponomareva (1975). (Sheard, 1965, 1967; Sebastian, 1966; Brinton, 1967a, 1973; Razouls and Thiriot, 1968; Roger, 1968c, d, 1974b, c, 1978a; Vives, 1968; Gopalakrishnan and Brinton, 1969; Guglielmo, 1969; Karedin, 1969b; O'Riordan, 1969; Antezana, 1970; Baker, 1970; Casanova, 1970, 1974; Casanova-Soulier, 1970a; James, 1970; Weigmann, 1970, 1974a, b ; Wiebe and D'Abramo, 1972; Beaudouin, 1973, 1974; Brinton and Gopalakrishnan, 1973; Casanova et at., 1973; De Decker, 1973; Talbot, 1974; Ponomareva, 1976c; Youngbluth, 1975; Taniguchi, 1976; Weigmann-Haass, 1976; McWilliam, 1977; Sipos, 1977; Rottman, 1978.)

Xtylocheiron maximum Hansen, 1908 Brinton, 1975: 218, figs 116, 124d.

Brinton (1975) has produced a revised map of its distribution (Fig. 14) that is different in several aspects from that in Mauchline and Fisher (1969). Atlantic Ocean : its distribution now extends northeastwards towards the Barents Sea; it occurs in the Gulf of Mexico but Brinton omits to show its occurrence in the western Mediterranean. He indicates its occurrence as far south in the Atlantic as 55"s. Pacific Ocean : Brinton shows it absent from the waters west of Baja California ;

60

40

20' 00

20' 40

60'

1200

160°

160°

1200

40°

40°

FIQ.14. Distribution of Stylocheiron maximum and S. robustum (after Brinton, 1975).

80°

3. DISTRIBUTION AND SYNONYMY

419

it occurs in the South China Sea. Indian Ocean: Brinton shows it present Bhroughout the, ocean between latitudes l0"N and nearly 60"s. (Sebastian, 1966; Brinton, 1967a, 1973, 1979; Roger, 1968c, d, 1974c, 19788,; Wiborg, 1968; Gopalakrishnan and Brinton, 1969; Baker, 1970; Casanova, 1970, 1974; James, 1970; Day, 1971; Franqueville, 1971; Wiebe and D'Abramo, 1972; Brinton and Gopalakrishnan, 1973; De Decker, 1973; Talbot, 1974; Youngbluth, 1975; Ponomareva, 1976c; Taniguchi, 1976; Lindley, 1977; McWilliam, 1977; Reid, 1977; Sipos, 1977; Robehon and Roberts, 1978; Rottmm, 1978.)

Stylochiron robusturn Brinton, 1962 Brinton, 1975: 218-220, figs 117, 124e.

The revised distribution given by Brinton (1976) is the same as that in Mauchline and Fisher (1969) for the Atlantic and Pacific Oceans but includes the records of James (1970) in the Gulf of Mexico (Fig. 14). It has also been found to occur in the southern Indian Ocean between latitudes 10"sand 30"s.(Youngbluth, 1975.)

CHAPTER 4

THE LARVAE The names used t o describe the larvae of euphausiids are nauplius, metanauplius, calyptopis and furcilia. These types of larvae and their nomenclature are discussed and defined in Mauchline and Fisher (1969). Williamson (1969) suggests that the terms nauplius, zoea, megalopa and juvenile be used but this is liable to cause confusion rather than simplification. Mauchline and Fisher (1 969) divide the development of the furciliae into phases as a means of classifying larvae. The larvae in each phase are then examined to determine the number of moults, that is larval stages, in each phase. The stages are numbered sequentially so detailing the number of moults in the developmental sequence. A similar scheme of stages is used by Endo and Komaki (1979) to describe the development of Thysanoessa Zongipes. Casanova ( 1974).proposes calling phases of development furciliae I t o 111; her system of presentation of the results of an analysis of the larval development of a species is useful in making comparisons with other species. Silas and Mathew (1977) review the various approaches used in studies of larval euphausiids. Mauchline and Fisher (1969) have reviewed the anatomy and histology of the male and female genital systems. Reference t o the androgenic gland described by Zerbib (1967) in Meganyctiphanes norvegica was omitted; this endocrine gland occurs close to the external opening of the vas deferens and is responsible for the differentiation of the primary and secondary sexual characteristics of the male. Hollingshead and Corey (1974) and Kulka and Corey (1978) have provided further figures of developmental stages of eggs within the ovary of M . norvegica and Thysanoessa inermis along with comparable figures of lobes of the testes. The ovaries a t different stages of maturation were dissected from the animals and photographed by Roger (19738).

Species in Group A of the genus Euphausia usually produce spermatophores in the left urn deferens, the lobes of the testes on the right side being markedly less developed (Mauchline and Fisher, 1979). Sebastian (1966) h d s that only one spermatophore is carried in Thysanopoda

4.

THELARVAE

421

tricuspidata whereas the normal two are carried by T . orientalis and Stylocheiron longicorne. The spermatophore is normally transferred by the male and attached to the thelycum of the female. It usually remains attached there for some time after the spermatozoa have discharged from it into the spermathecum and is probably not lost by the female until she moults. The four Euphausia species in the “gibba” group, namely E . hemigibba, E. pseudogibba, E. paragibba and E. gibba, appear to be exceptional in that the discharged spermatophores are not retained by the females. James (1977) believes that males of the latter three species transfer the spermatophores briefly to the structural elements of the thelycum present in the seventh thoracic segment of the females so that the spermatozoa can discharge into the spermathecum. The thelycum of E. hemigibba is the least specialized and in this species the male may hold the spermatophore briefly on or close to the spermathecum t o effect discharge of the spermatozoa. Brinton (1978) reports nine occurrences of spermatophores attached to the first pair of pleopods, one occurrence on the second pair of pleopods, and three instances of attachment to the gills of males. The species involved were Thysanopoda aequalis, Euphausia pacijica, E . similis, Thysanoessa gregaria, Nemtoscelis dificilis and N. microps but these were extremely rare occurrences. The seasonal changes in the percentage of males and females in populations of Meganyctiphunes norvegica and Thysanoksa raschi carrying spermatophores is shown in Mauchline and Fisher (1969). Transference of the spermatophores from the males to the females in a population takes place some considerable time before egg laying. Wiborg (1971), examining populations of Meganyctiphanes mrvegica in the Byfjord and Hardangerfjord in Norway, and Hollingshead and Corey (1974), examining populations of the same species in Passamaquoddy Bay in eastern Canada, confirm that transference takes place over a period of about three months. Similar results were obtained by Kulka and Corey (1978) in a study of Thysanoksa inermis. Berkes’ (1976) data are insufficient to draw any conclusions about the length of the transfer period. It is not known why spermatophores should be transferred to the females when the ovaries are still in an early stage of maturation. Mauchline (1972), in considering the breeding strategies of oceanic and deep sea pelagic organisms, suggests that early mating may be necessary for the initiation of the final stages of ovarian maturation. It is not a single act of mating that is being discussed here because the females are usually actively growing in body size and consequently they are moulting and have t o be remated. The frequency of moulting is once every two or three days up to

422

THE BIOLOGY O F EUPHAUSIlDS

about once every twenty or thirty days, depending upon the species, its body size and the environment. Thus, there is opportunity for complex working of endocrine systems stimulated by the mating processes. Brinton (1978) points out that fertilization is always accomplished by one spermatophore in Euphausia pacijka. This also seems to be true in Meganyctiphanes norvegica. ThysanoGsa species are fertilized by either one or two spermatophores while Brinton states that Nematoscelis species appear to require two spermatophores. Examples of the ranges in sizes of the eggs of different species are given in Mauchline and Fisher (1969). Ponomareva (1969) found that eggs of Euphmusia diomedeae from' the Indian Ocean ranged in diameter from 0-30 t o 0.42 mm with a perivitelline space of 0.04 mm. She obtained large eggs, 0-71-0-87 mm in diameter, which she considers are those of Thysanopoda tricuspidata. Ponomareva (1969) examined the development time of the eggs of Ewphausia diomedeae and found that they require 16 h to develop from the stage of cleavage into two blastomeres to a fully developed nauplius. The series of early larval stages in species that lay their eggs freely in the sea consists of a nauplius, that emerges from the egg, a subsequent second nauplius, a metanauplius and three calyptopes. Species in some genera carry their eggs attached to the thoracic legs; these eggs hatch a t a later stage, the second nauplius, which develops into a metanauplius. The abdomen and stalked eyes develop in the three successive calyptopes. All species of euphausiids pass through this developmental sequence of stages (Mauchline and Fisher, 1969).

Table I in Mauchline and Fisher (1969) contains measurements of the early larval stages of 36 species of euphausiids and lists the relevant literature. Detailed descriptions of the individual larvae can be obtained from that literature. Additional information is now available on 14 of these species and is presented in Table I11 along with data on a further 13 species. Consequently, information on the early development of 49 species has now been published. Le Roux (1976) has demonstrated marked annual variations in the body sizes of the three calyptopes of Meganyctiphanes norvegica and Nyctiphanes couchi in the Gulf of Morbihan, France. This is probably a general feature of larvae of most species and so the measurements given in Mauchline and Fisher (1969) and in Table 111 are presented primarily as ranges in size. Mathew (197 I ) describes the larvae of Euphausia distinguenda from the northern Indian Ocean but Brinton (1975) has re-named these populations E . sibogae; Bhe name E . sibogae is used in Table 111.

TABLE111. RANGEIN S m (mm) OF EARLYLARV~LL STAGES

Naupliua Speciea Thysanopoda T . rnonacantha T . tricwpidata T . a.eqwlis T . pectinata; Meganyctiphanes M . norvegica Nyctiphanw N . couchi Euphausia E. krohni E . brevis E. diontedeae E . crystallorophias E . sibogae E . fallax E. gibboidw E . sanzoi E . hmnigibba E . hanseni ThysanoZsaa T . gregaria T . longipea

I1

Metanauplius

I

,I1

1.00-1.1 2

1.36-1.50

1-90-2.16 1.20-1.44

2.90 2-50-2.74 1.70-1.90

0.48

0.50-0.52

(1.03)

0.56

0-57

0*80-1*00

0.40-0.60

0.90-1.40 0.88-1 *03 1.161.23 1.35-1.50 0.96-1.15 1.17-1.35 1.09-1.27 1.01-1.23

I

0.48

0.48-0.53

Calyptopes

0.51-0.56

0.53-0.6 1

(1.59)

III 3.40 2.90-3.40 2.20-2.30 4-00 (2-40)

References

7 1, 5 6, 7 7 1, 6, 11

1.38-1.44

1* 80-2.20

1, 4, 6, 11

1.38-2.00

1, 6 1, 6

1.00-1.40

1.50-1.70 2.25-2-80 1.49-1.58 1.80-2.14 1366-1.98 1'68- 1'92 1.60 1.70-2.10

1.80-2.80 1.70-2.40 2.14-2.28 3.66-4.45 1.9%2*00 2.48-3.07 2.34-2.7 1 2.18-2.79 2.00-2-60 2.90-3.60

6 13

0.96-1.30

1.40- 1.80 1.43-1'93

2.00-2.50 2.28-3.20

1, 6 15

1'20-1.60

2, 7 12 2 14 9

10

contd.

TABLEIII-contd. Nauplius Species Nematoscelis N. di&ilis N. megalops N. tenella N. microps N. atlanticu N. gracilis Stylocheiron S. carinatum S. suhrni S. longicorne S. abbreviatum S. maximum

I

0.43-0.53

Calyptopes

I1

0.60 0.53-0.57

0.90-0.96

Metanauplius

0.80

I

XI

Ill

1.30-1.60 0.80-1.70 1.20-1-30 1.30-1-40 1.40-1.50 1.30-1.40

1.80-2-10 1.60-2.50 1.60-1.90 1.80-2.10 2.1Cb2.30 2-00-2-20

2.00-2.80 2.20-3.30 2.40-2-70 2-60-2.80 2.60-2.90 2-50-2.70

1.60-1’75 1.20-1.28 1-30 1.80-2.24 2.47-2.70

1.80-1.96 1.70-1.76 1.70-1-90 2.66-2,75 3.92(?)

2.08-2.59 2.00-2.50 2.30-2.40 2.40-2.87 3.85

References

1, 8 1, 6, 8 1, 8 1, 8 8 8 1, 3, 7 1, 6

6 1, 6, 7 1, 4, 6

Mean sizes are shown in brackets, single measurements not in brackets are of doubtful accuracy. This is additional information to that of Table I in Mauchline and Fisher (1969). References: 1, Mauchline and Fisher, 1969; 2, Mathew, 1971, 1975; 3, Mathew, 1972; 4, Casanova, 1972; 5, Knight, 1973; 6, Casanova, 1974; 7, Brinton, 1975; 8, Gopalekrishnan, 1975; 9, Knight, 1975; 10, Knight, 1976; 11, Le Roux, 1976; 12, Pertzova, 1976; 13, WeigmannHaass, 1977; 14, Knight, 1978; 15, Endo and Komaki, 1979.

4.

425

THELARVAE

The third calyptopis moults to the first furcilia. This larva has the eyes free of the carapace and is a miniature adult in body form but lacks most of the thoracic legs and pleopods. The successive furciliae are recognized principally through the developmental sequence of the pleopods and abdominal photophores and by the reduction in number of the terminal spines on the telson; there are seven spines in the fist furcilia reducing to one in the adolescent (Mauchline and Fisher, 1969). A pleopod develops first as a non-setose rudiment which becomes setose at the following moult. Anterior pairs of pleopods develop before posterior pairs. Some species follow a strict developmental pathway, e.g. Thysanopoda acutifrons has a furcilia I with hree pairs of rudimentary pleopods (denoted 3’), a furcilia I1 with t ree pairs of setose and two pairs of rudimentary pleopods (denoted 3” 2‘) and a furcilia I11 with five pairs of setose pleopods (denoted 5”). This course of development is written as follows :

k

3’

+ 3” 2’

d

5’‘

Other species have a dominant pathway along which the majority of larvae develop but variations in the development are common (Mauchline and Fisher, 1969). For example, in Meganyctiphanes norvegica most larvae follow the dominant pathway: 3’

+ 3‘’ 2‘

d

5’’

but a notable number, varying between populations and seasons, will follow one of the following: 0’

+

4’

1’ + 1”2‘

_?,

4” 1’

4

3“ 2’

+ 5”

5“

1’ + 1”3‘ + 4” P’ + 5” 1’ ---+

1’’ 4’ + 5”

2’

+ 2” 3‘ +

2’

+ 2“ 2‘ + 4“ 1’ + 5”

3’ + 3” 1’

4’

__+

4” 1’

5”

-

__f

4’’ 1‘

5”

5”

The forms of early furciliae found in some fifty species are described by Mauchline and Fisher (1969). Additional information is now available on 32 of these species and the development of a further twelve

426

THE BIOLOGY OF EUPHAUSIIDS

species has been described. All the available information is summarized in Figs 15 and 16. Mathew's (1971) Ewphawia distinguenda has been renamed E . sibogae (see Brinton, 1975). The development of E . recurva described by Sheard and shown in Mauchline and Fisher, Fig. '47, is not that of E . recurva; the larvae described by Sheard belong to another unidentified Euphausia species according to Brinton (1975).

Mauchline and Fisher (1969) discuss a t some length the reasons why developmental pathways may be variable in some species and more or less constant in others. They point out that the dominant pathway of development of a species not only varies between different sea areas but can also vary in the same sea area at different times. This has been demonstrated recently in furciliae of E . pacifica by Endo and Komaki (1979). The developmental pathway is probably influenced by the availability of food but other environmental conditions are no doubt involved. Le Roux (1973a, b ; 1974) has examined the influence of food, environmental temperature and removal of the eyes on the larval development of Meganyctiphanes norvegica and Nyctiphanes couchi. These are difficult experiments t o perform satisfactorily in the laboratory because the general conditions of culturing are themselves likely t o affect the developmental sequence. Zelikman (1968) and Makarov (1971b; 1974a) discuss in full the significance of dominance among larval forms of a species. They stress the point that variant forms are not the result of accidental happenings within the development of a larva but are undoubtedly a direct reflection of the environmental conditions in which the larva is living. Changing environmental conditions, over a more or less extended breeding season, cause a change in the dominant pathway of development; or they may cause the appearance of other pathways of development where only one existed before. A taxonomic group of species probably has a dominant pathway of development which is optimal for these species (Figs 15 and 16) and is used under optimal or near optimal environmental conditions for these species. When environmental FIG.15. Development of the early furciliae of species in the genera Thpanopoda, Meganyctiphanes. Nyctiphanes, Peeudeuphausia and Euphawia. The blocks are representative of the relative numbers of each type of larva recorded in any one species. Open squares represent most probable types of larvae but not s d c i e n t evidence is available to be categorical. Question marks are doubtful records. (1) Mauchline and Fisher, 1969; (2) Jones, 1969; (3) Ponomareva, 1969; (4) Mathew, 1971; ( 6 )Mathew, 1972; (6) Gopalakrishnan, 1973; (7) Le Roux, 19738; (8) Casanova, 1974; (9) Le Roux, 1974; (10)Makarov,197413; (ll)Talbot, 1974; (12)Brinton, 1975; (13)Gopalakrishnan, 1975; (14) Knight, 1975; (15) Pertzova, 1976; (16) Weigmann-Haaas, 1977; (17) Knight, 1976; (18) Le ROUX,1976; (19) Knight, 1978; (20) Endo and Komaki, 1979.

-0

THYSANOPODA T monacantha T tricuspidoto 7:aequalis T microphthatma T acutifrons T pmtihato T orientatis T cristata I: egregia 7: mrnuta/spinicaudata

--

-@J

-*

-i

-0

I

I

MEGANYCJtPHANES Ynorvegica NYCTIPHAN€S N oustratis N capensis N couchi N simplex PSEUDEUPHAUSIAI I? totifrons

I? sinica EUPHAUSIA E krohni E mutico E bre vis E diotnedeae E recurva E superba E vattentini E hens E frigida E pocifica E nona E crystaltorophias E tenera E sitnilis E sibogae E gibboides E fatlax E sonzoi E pseudqibba E hemigibba E spinifera E honseni E longirostris E triocontho

1 1 1,12 1,12 1

I :

I

-

I

1,8,9,18

_ I

I-' I

I -

I

--I

I

1

I I

1,12 1

I

1,12 4 12,14 12,19 12,17 12

-n

428

NEMATOSCEL./s N difftciks N megolops N tenella N microps N.otlontica nl. lobato nl.gracilis

srrmxmoiv S mrinatum S . offine S suhmi S efongatm S,longicorne S obbreviatwn S maximum

THE BIOLOGY OF EOPHAUSIIDS

-a

m

I

I I I

I

I1,6,13 1,8,13 1,13 1,13 1,13 1.13 1,13

I

I I

FIQ. 16. Development of the early furcilia6 in species in the genera Thysanokkaa, Nenzatoscelia and Stylocheiron. Format of figure and references as in Fig. 15.

conditions change, for example, later in the breeding season, and become less optimal for the species, the pathway of development changes in response to these conditions. This “variant” pathway is probably not optimal for the species except possibly under the sub-optimal environmental conditions., Consequently in studying the development of a species sampling should be carried out, not only throughout the geographical range of the species, but also throughout the seasonal period of occurrence of the larvae. The development of the later furciliae is referred to in Mauchline and Fisher ( 1969) and in the papers cited in Figs 15 end 16. Marr (1962) demonstrated an ontogenetic migration of the larvae of Euphausia superba and Mauchline and Fisher (1969) discuss the vertical distributions of euphausiid larvae and their relationships to such migrations and to diurnal vertical migrations performed by some of the later furciliae. Hempel and Hempel (1978) and Nast (1979) examine bhe distribution of larval E . superba in the Atlantic sector

4.

THELARVAE

429

of the Antarctic. Few eggs and no nauplii were taken in the upper 150m of the water column. 'There was evidence of some vertical migration of furciliae I, I1 and I11 into the neuston a t night. Haury (1976a) found that the modal depth of occurrence of euphausiid eggs in the California Current is at about 100 m but he did not identify the species. ,Eggs, probably those of Thysanoessa raschi, were concentrated in the surface 60 m of a water column of total depth 170 m in a northern Norwegian fjord (Hopkins and Gulliksen, 1978). Makarov (197%) found that the eggs, nauplii and metanauplii of Euphausia frigida, E . triacantha and Thysanohsa species occurred deep and that the larvae of these species performed an ontogenetic migration in the sea area off north-eastern South Georgia. The depth at which several species lay their eggs is still unknown. The investigations of populations of the Antarctic Euphusia superba have been relatively inconclusive (Hempel, 1979), although this species, like other epipelagic and shallow living species, probably lays them close t o the sea surface. Information on the depths at which eggs are laid by the bathypelagic Bentheuphausia amblyops and species in the genus Thysanopoda is not available. All euphausiid eggs so far examined are slightly denser than sea water and so sink. The furciliae, however, are primarily phytophagous and live in the surface layers. There will be some advantage in restricting the extent of the ontogenetic migration required to be performed by the earlier larvae, which probably do not feed to any great extent, as discussed below. The only way of doing this is by the females laying the eggs high in the water column so that the nauplii hatch out a t relatively shallow depths. The food of larval euphausiids has received little attention (Mauchline and Fisher, 1969).The mouthparts are not functional in the nauplii and metanauplius but become functional in the first calyptopis. Pavillon (1978) examines the capability of eggs, nauplii, metanauplii and first calyptopes of Euphausia krohni to absorb organic nutrients directly from the sea water. He introduced the following a t concentrations likely t o be encountered in the natural environment : glycine, alanine, serine, cysteine, asparbic acid, glutamic acid, methionine, byrosine, thiamin, riboflavin, pyridoxin, cobelmin and palmitic acid. Amino acids were absorbed in greater quantities by larvae than by eggs. Little cobalamin and pyridoxin was absorbed by the eggs but these were absorbed appreciably by the larvae. The absorption of riboflavin increased with time in both eggs and larvae.

CHAPTER 5

VERTICAL DISTRIBUTION AND MIGRATION Vinogradov (1970) has reviewed the vertical distribution of oceanic zooplankton, placing the euphausiids in the context of the oceanic biomass. The primary information available in his book, and not in Mauchline and Fisher (1969), is the vertical distribution of the biomass of euphausiids as a whole. Samples were taken to depths greater than 6000 m but no euphausiids were caught at depths greater than 6000 m. He recorded the following changes in euphausiid biomass with depth in the subarctic region of the Pacific Ocean: 0-50m, 14.8 mg/m3; 50-500 m, 4.6 decreasing to 2.1 mg/m3; 500-1500 m, 0.2 mg/m3; 15002500 m, 0.02-0.06 mg/m3; 2500-3000 m, 0.70 mg/m3; 3000-5000 m, 0-0-03 mg/m3. These biomasses represent 1-7 % of the tot.al biomass of plankton in the surface 500 m layer and less than 1% in the deeper layers. Biomasses of euphausiids in the tropical regions of the Pacific Ocean were less than in the subarctic regions but represented 2.413.4% of the total plankton biomass in the 400 m water column. Euphausiids are pelagic organisms, the majorihy of the 85 species living in the epipelagic or shelf environments. Nineteen species are classified as mesopelagic or bathypelagic (Table IV). Many species me known to perform diurnal vertical migrations, rising towards the surface at night and descending to deeper layers during the day. Mauchline and Fisher (1969) reviewed the available information on the vertical migrations of 39 species. New information has been obtained on the migratory behaviour of many of these species as well as on that of a further 26 species (Table IV). The bathypelagic species Bentheuphausia amblyops does not perform a diurnal vertical migration and this is probably true of the other bathypelagic species listed in Table IV. Fourteen species are conaidered to be mesopelagic and five of these have been shown to perform a diurnal migration, certainly in some sea areas on some occasions. The remaining nine species either do not migrate or only move through a small vertical range. Several epipelagic Stylocheiron species have been shown not t o perform a migration (Table IV). All other epipelagic euphausiids investigated, wibh the possible exception of

Species

Distributional WPe

M= migrates

Bentheuphawia amblyops Thysanopoda mcmmantha

Bathypelagic Mesopelagic

No M

T . cristata T. t r i c u a m T . aequalis T . astylata

Mesopelagic

(m)

Average night depth

(m)

> 1000 >800-400

700-100 >900-100

300

-

600-300 >800-200 400-0 700-0

400 300

S

700-500

400-0

M

Below

600-300

300-100

Mesopelagic

M

S(?)

70&600

300-0

Mesopelagic Bathypelagic

NO

Coastal Coastal Coastal

(m)

Below

M

S M No No No M M M M

Diataw migrated

-

>800-400 700-400

Bathypelagic Bathypelagic Bathypelagio

Total vertical range

(4

S

T . obtwifrons

T . orientalis T . mkrophthalnta T . acutqrons T . cornuta T . egregia T . spinicaudata Meganyctiphanes norvegica Nyctiphianerr couchi N . australis N . capenaris

(8)

Average day depth

M

I

T . pectinata

Species reaching surface atnight

500 100

-

S S S

S

700-500

> 500

>900-0

1000-100

400

700-500 >800-0

> 500

900-700 500-200 >loo0 >loo0 >loo0 400-100 100-0 <200-10 <100-0 200-100 100-0 50-10 3&0

>1000-100

-

1500-0 >400-0 400-0 >90-0

References 1, 31 5, 11, 17, 20, 30, 31 11, 17, 31 5 , 11, 17, 31

2, 5, 8, 9, 14, 17, 20, 31 5, 8, 16, 20, 31 11, 17, 20, 31 11, 17, 31 8

11, 30,

17, 30,

300-400 5 1 1 1 200 5,9,12,14,23
-

TABLEIV-contd.

Species

Distributional tYP

Species reaching surface M = at night (S) mipates

Average day depth (m)

Average night depth (m)

Total vertical range (m)

Distance migrated

150 50

(m)

M M

S S

300-50 80-50

100-0 80-0

>500-0

M

S S S S

100-0 150-0 200-0 100-0 50

>900-0

M M M

700-500 700-150 600-400 600-300 400-200

E . diornedeae

M

S

200-0

>200-100

100

E . recur-va E . auperba

M M

S S

300-0 70-0

700-0 >900-0

300 50

E . vallentini E . lucena E.frigida E . paci&a

M M M

S

S S

>lo0 <50 500-250 100-50 400-150 100-0

M

S

> 300

M

S

N . simplex Pseudeuphausia latijrons P . Binica Euphausia americana E. exhia E . krohni E . mutica E. breh

E . nana E . crystallorophias E . tenera E . Sirnilis

Coastal Coastal Coastal

> loo? 700-150 100-10

500-300

100-0

700-0 >900-0

>900-0

400 300 <200 200 300

100

> 750-0 >400-0

200

>900-0

250

150-0 100-0

References 2, 25 5 5 1 5, 8, 9, 14, 21 5, 20, 27 2, 3, 5, 8, 9, 14, 16, 20 3, 10, 11, 18, 31 1, 27 5, 6, 15, 24, 28, 34, 35

22 5 2, 5, 7, 13, 19, 27, 33 1

5, 11, 17, 20, 31

E . similis var. armah E . mucronata E . aibogae E . distinguenda E . lantelligera E’. gibba E. gibboides E . fallax E. sanzoi E . pseudogibba E. paragibba E . hmnigibba

E . spinifera E . hanseni E . longirostris E . trimantha Tessarabrachion oculatunz Thysanoiissa spinifera T . longipes T . inspinata T . inemis T . longimudata T . parva T . gregaria T . vicinu T.m m r a T . raschi Nematoscelis d i f i i l i s

Mesopelagic

Mesopelagic

M M

S S

300-0 300-0 1500-5OOP 1000-O?

5009

M M M

S 50 S

600-400 500-200 400-300

200-100 900-0 300-50 ~ 9 0 0 - 5 0 400-0 200-100

200 200 200

5, 8, 27

M M M

S S S

100-25

25-0 100-0 100-50

>200-0

50

3

>900-0

300

11 2, 5, 8, 9, 14, 16

M M M M M M M i NO MP

S S S S S S 3

750-0

200 200 50 150 100 200 300

M Coastal

400

600-400

500-250

> 200 100-0 250-50 400-50 300-0 400-0

> 500

250-0

> 400 100-70 150-0 150-0 100-0 10&0

> 500

S

600-400

500-100

S

100-50

100-0

300-50

100-0 300-0

M

S

M

S

> 300

> 400-200 100-0 300-0 300-0 400-0 400-0

> 500 100&0

1 6

17, 20 17, 31

5

7, 27 13, 27 7, 27 7 5, 12, 23 5 , 23, 32 8

250

2, 5 , 8, 9, 14, 21, 26, 27

>750-0

50

5

>200-0 600-0

100

5, 12, 23, 29 2, 27

P

SpfXie'8 reaching Speck

Distributional type

M= migrata

Burface at night

(S)

depth (m)

Average night depth (m)

Total vertid range (m) 2500-100

Average

h Y

Distance migrated

(4

N . mgdops

Mesopelagic

M?

> 50

500-300

300-100

N . tenella

Mesopelagic

M

100

600-300

400-100 >900-200

300

N . micropa

M

Below

800-400

400-200 >900-100

300

N . atlantica

M

Below

600-400

400-100

200

N . gracilis

M

S?

600-500

500-200

Nematobrachion flea%?

M

50

600-400

500-300 >600-50

100 100

700-50

200P

References 5, 8, 9, 14, 21, 26 2, 5, 8, 11, 16, 17, 20, 31 5, 8, 10, 11, 16, 17, 20, 31 2, 14, 17, 20, 31

N . lobata

N . sexapinomm

Mesopelagic

M

100

600-400

500-300 >600-100

N . boopis

Mesopelagic

No

100

600-500

600-500 > 1000-250

Stylocheiron carinaturn

M?

S

300-100

200-0

>goo-0

S . aBne

No

S

300-200

300-0

800-0

loo?

3, 10, 11, 17, 20, 31 11, 17, 20, 27, 31 3, 17, 20, 30, 31 5 , 8, 11, 17, 20, 31 2, 5, 8, 11, 17, 20, 31 8, 11, 17, 20, 27, 31

S . suhmi

No

S

200-0

200-0

S. microphthalma S. inaubre S. elongatum

No

S

200-0

200-0

Mesopelagic

No

200

500-300

500-200

>900-200

S. indicum S. longicorne

Mesopelagic

No

300-100

300-100

400-100

300-0 2000-50

?

S

400-200

400-100

2000-50

300-150 45&300

500-200

8. abbre viaturn

s.madmum S. robusturn

Mesopelagic Mesopelagic

.

? ?

150 300

> 450

>900-0

2, 5, 8, 17, 20, 31 17, 31

loo?

5 , 8, 11, 16, 17, 20, 31 4 2, 5 , 8, 9, 11, 14, 17, 20, 27,

31 2, 5, 8, 9, 11, 14, 17, 20, 31 8,9, 11, 20,27 20

References: 1, Brinton, 1962; 2, Brinton, 196713; 3, Naumov and Ponomareva, 1967; 4, Silas and Mathew, 1967; 5, Mauchline and Fisher, 1969; 6, Shevtsov and Makarov, 1969; 7, Taniguchi, 1969; 8, Baker, 1970; 9, Casanova, 1970; 10, Ponomareva, 1971; 11, Roger, 1971; 12, Wiborg, 1971; 13, Alton and Blackburn, 1972; 14, Casanova, 1974; 15, Pavlov, 1974; 16, Roe, 1974; 17, Roger, 197413; 18, Weigmann, 1974s; 19, Marlowe and Miller, 1975; 20, Youngbluth, 1975; 21, Andreu, 1976; 22, Bartle, 1976; 23, Berkes, 1976;, 24, Fischer, 1976; 25, Kanaeva and Pavlov, 1976; 26, Weigrnann-Haass, 1976; 27, Youngbluth, 1976; 28, Everson, 1977; 29, Hopkins and Gulliksen, 1978; 30, Hu, 1978; 31, Roger, 19788; 32, Sameoto, 1978; 33, Greenlaw, 1979; 34, M o b , 1978; 35, Nest, 1979.

436

THE BIOLOGY O F EUPHAUSIIDS

Thysanoessa gregaria, appear t o perform more or less regular diurnal vertical migrations. Demonstration of a diurnal vertical migration of a species is sometimes inconclusive because some species appear t o migrate irregularly, A regular migration produces larger catches at night in the surface layers. Sampling methods have to be examined because avoidance of and escapement from towed nets may affect the results (Zhuravlev, 1976b). Taken to the extreme, avoidance of and escapement from nets in the surface layers during daylight followed by more efficient capture during the hours of darkness could produce results simulating a diurnal vertical migration. Reports of the capability of euphausiids to avoid capture by nets are frequently contradictory (Mauchline and Fisher, 1969). There are three primary variables in samplingdimensions of the net and the duration and the speed of towing. Mesh size of the net material is usually, to some extent, dependent upon the designed size and speed of towing of the net. A useful introduction to sampling methods is contained in Anonymous (1968). Special studies on the behaviour of euphausiids in relation to different samplers are relatively few (Mohr, 1978). Aron and Collard (1969) found that the three species Euphausia pacijica, E . hemigibba and Nematoscelis dificilis were caught with the same efficiency by a 6-ft (1.83 m) Isaacs-Kidd midwater trawl towed at two speeds, about 1-0 and 2.1 m/s. The Isaacs-Kidd trawls, the 6-ft (1-83m) and 10-ft (3.05 m ) versions (Friedl, 1971b), are frequently used for capturing euphausiids because these nets are larger than the majority of plankton nets; their large sizes are considered to reduce the proportion of animals avoiding capture, although Allen (1972) used 1.0 and 0.5 m Clarke-Bumpus nets and detected no significant escapement or avoidance of these small nets, even during the day. The mesh sizes or combinations of mesh sizes along the length of Isaacs-Kidd trawls vary between different investigations. Estimations of capture efficiency and degree of escapement through various cod-ends with the different mesh sizes have been studied by Michel and Grandperrin (1970). They found that escapement through a 4 mm mesh was a function of the duration of the tow and the vertical distribution of the euphausiids. Wiebe ( 1972) found that increased length (duration) of tow reduced the sampling errors to a greater extent than changing to a larger net filtering a larger volume of water in a shorter towing distance. Angel (1977) has summarized many of the current problems involved in collecting and interpreting samples of oceanic pelagic organisms. It is inadvisable to use the same design of nets t o sample eggs and

5. V E R T I C U DISTRIBUTION AND MIGRATION

437

larvae on the one hand and the larger more active adults on the other. Quantitative comparisons of data from different nets are often desirable and were attempted by Roger (1968a, b) for an Isaacs-Kidd midwater trawl and a 1 m plankton net. Several new n e b are available including the Rectangular Midwater Trawl (RMT) combination net with a 1,elemeteringsystem for opening and closing it a t known fishing depths (Baker et al., 1973; Pommeranz, 1978) and the modified Tucker trawl that can be closed by messenger (Hopkins et al., 1973). Pearcy et al. (1977) have described a modified Isaacs-Kidd midwater trawl that incorporates a 1 m2 multiiple plankton sampler with five codends. The Longhursf-Hardy plankton recorder is increasingly being used to sample euphausiids, especially larval stages. Haury (1973) has examined the problem of “hang-up” of organisms in the collector net in front of the recorder. Some animals are known t o enter the collector net and remain there for an indeterminate period before being washed down into the recorder. This delay or “hang-up” is variable but can be minimized by careful design of the collector net. Johannessen ( 1977) found that Meganyctiphunes norvegica and euphausiid larvae showed no signs of hang-up in the test samples of plankton taken by h* in the Korsfjorden, Norway. The critical design parameters of this equipment are discussed by Haury et al. (1976). Interesting and potentially useful sampling devices are the suction tube for midwater organisms described by Croce and Chiarabini (1971) and the light trap for plankton described by Jones (1971). A diving saucer was used by Franqueville (1970) Qo make visual observations on the distribution of pelagic organisms, including euphausiids. Results obtained from the saucer were compared with data obtained from IsaacsKidd midwater trawls taken at the same time. Difficulty was experienced in identifying the euphausiids visually from the saucer. Mauchline (unpublished) found ehat the lights of the Vickers Pisces submersible seemed to attract euphausiids and other organisms, thus distorting estimates of density. Mauchline and Fisher (1969) discuss the aggregation of euphausiids into patches, shoals and surface swarms or rafks. Presence of such aggregations seriously affects the interpretation of quantitative samples taken to assess the vertical distributions and migrations of species. New information on aggregations of euphausiids is given in Chapter 11. There are a number of factors that modify the general pattern of the diurnal vertical migration of a species. Larvae often behave differently from the post-larval stages, usually occurring closer to the surface at all times and frequently being members of the neuston

438

TEE BIOLOGY OF EUPHAUSIIDS

at night. Kanaeva and Pavlov (1976) found that larvae of Nyctiphanee simplex off Peru migmted to the surface in the morning and again in the evening, while juveniles and adults only occurred at the surface in the evening. The age or body size of an individual of a species can determine how closely it will approach the surface at night. Larger individuals of a species tend to live at greater depths than smaller individuals and this vertical gradient of size can even be maintained within the population throughout a diurnal vertical migration. This has been shown to occur in Meganyctiphanes norvegica, Euphawia krohni, E . pacijica, E . triacanthu, Thyeano&sa longipes and T . inermig (Mauchline and Fisher, 1969). It also occurred in populations of Meganyctiphunes norvegica in the Byfjord, Norway (Wiborg, 1971). Roger (1971) found evidence of a' gradient of increasing body size with increasing depth of occurrence in populations of the following species in the equatorial Pacific Ocean: Thysanopoda orientalis, T . monacantha, T . pectinata, Euphuusia paragibba, Nematoscelis microps, N . gracilis, Nematobrachion boopis, Stylocheiron abbreviatum and S . maximum. Gravid females may be an exception in that they have been found in the surface layers during both day and night. No such relationship was found within the population of T. rawhi in the Clyde sea area (Mauchline and Fisher, 1969). Ponomareva found a very different situation in populations of this species in the Barents and Okhotsk Seas where smaller individuals lived at greabr depths than larger individuals; this observation has not been repeated. Baker (1970) and Angel and Fasham (1973) andysed the vertical structure of the euphausiid assemblage off Fuertaventura, Canary Islands. There was a tendency for adolescents of most species to live at shallower depths than the corresponding adults. The patterns of vertical migration of the adolescents and corresponding adults were different. For example, adolescent Nematoecelis tenella did not perform a vertical migration while the adults did; in other species, both the adolescents and adults migrated diurnally but they had different depth distributions. Rudjakov and Tseitlin (1976) found that species of larger body size tended to perform more intensive migrations than species of smaller body size. The effects of the physical4hemical parameters of the environment, such as gradients of salinity, density, viscosity, temperature and pressure, are discussed briefly and inconclusively in Mauchline and Fisher (1969). Little new information is available (Youngbluth, 1976). The presence of a thermocline appears to restrict the upward migration of some species but not of others. The restriction may result from an increased density of food organisms associated with

5.

VERTICAL DISTRIBUTI~NAND MIGRATION

439

the thermocline, rather than t,he thermocline itself acting as a physical barrier (Mauchline and Fisher, 1969; Roger, 1971). Light is probably the dominant factor controlling the diurnal vertical migrations (Mauchline and Fisher, 1969 ; Roe, 1974). Roger (1947~)demonstrated the effect of moonlight on the night levels of occurrence of several species of euphausiids. He found that the vertical range of distribution of the species in the 0-200 m layer was shifted deeper, the degree of shift being almost directly proportional to the intensity of the moonlight. There were, however, differences between the vertical distributions of bhe species during the New moon and Full moon phases. These differences suggest the presence of an endogenous rhythm as part of the control system of a diurnal vertical migration. The effect of a total solar eclipse on the vertical distributions of euphausiids was reported by Bright et al. (1972). Samples were collected a t the surface and a t 25m depth in the Gulf of Mexico on 6-8 March, 1970. Four species, Euphausia americana, E . brevis, E . mutica and E . tenera lived deeper than the 25 m net during the day and only occurred in samples a t night. A fifth species, Stylocheiron carinatum, occurred in the 25 m net, but not the surface net during the day. It reacted to the eclipse, which took place close to noon, by migrating t o the surface and being completely absent from the 25 m net. The vertical migration of all these species on the night following the eclipse was different from that on the night before the eclipse. Ferrari (1971) suggests that the eclipse upset the endogenous rhythm controlling the vertical migration. The association of populations of euphausiids with isolumes means that they tend to live within a more or less restricted vertical layer of the water column. Cushing and Richardson (1956) produced evidence that such a layer of Nyctiphanes couchi in the North Sea was responsible for the traces recorded on an echo sounder. Evidence suggesting that euphausiids are sometimes responsible for some of the deep scattering layers (DSLs) recorded by echo sounders is reviewed in Mauchline and Fisher (1969). Two symposia (Farquhar, 1971; Andersen and Zahuranec, 1977) have stimulated work in this field. A series of papers have investigahed the role of euphausiids as sound scatterers in the oceans: Kinzer (1967, 1969a, 1971); Bary and Pieper (1971); Beamish (1971a, b ) ; Bradbury et al. (1971); Ebeling et al. (1971); Fried1 (1971a); Pearcy and Mesecar (1971); Pedenovi and Croce (1971); Sameoto (1972, 1976a); Farquhar (1977); Freytag (1977);Mohr and Fischer (1977); Pieper (1977); Reid (1977); Greenlaw (1979). According to Farquhar (1977), euphausiids have been shown

440

THE BIOLOGY OF EUPHAUSIIDS

to cause sound scattering at echo-sounder frequencies of 42, 107 and 200 kHz but not in the 1-20'kHz range where mesopelagic fish with gas-filled swim bladders are the dominant scatterers. A special study of acoustic scattering by Euphausia pacijica was made by Beamish (1971a, b) using a 102 kHz echo sounder and by Greenlaw (1977, 1979) and Steinberg and Mentjes (1978) using a wide range of frequencies. Beamish defined the back scattering cross section of a typical euphausiid at 102 kHz to be 1.4 x cm2. Sameoto (1972, 1976a), using a 120 kHz echo sounder, studied the relationship between the distributions of Meganyctiphanes norvegica, Thysanoessa raschi and T . inermis, on the one hand, and of the phytoplankton on the other. He relied on conventional net sampling to determine the densities of euphausiids in the scattering layers detected with the echo sounder. Hopkins et al. (1978) used a 120 kHz echo sounder to study the zooplankton sound scattering in the Balsfjord, northern Norway. The euphausiids Megunye~iphanes norvegica, Thysanoessa rasehi and T . inermis were partially responsible for sound scattering. The scattering layers were shown to be dynamic assemblages of organisms, their species composition changing with time on a diurnal and probably seasonal basis. Further investigations of the back scattering cross sections of euphausiids and their signatures on echo sounders may enable surveys of the type carried out by Sameoto to be made without recourse to net sampling except for confirmatory purposes. Mauchline and Fisher (1969) discuss possible reasons that would cause an evolutionary development of a diurnal vertical migration. They concluded that such a migration would probably confer a range of benefits on an individual and species rather than one major advantage, such as increased availability of food. Baker (1970) found evidence suggesting that, within a genus, there is a marked tendency for vertical separation of non-migrant species whereas overlapping of vertical ranges may be present in migrating species. Analysing the same data further, Angel and Fasham (1974)point out that the vertical distributions of the adolescents and adults of the different species, whose ranges of body size vary, may be indicative of resource partitioning. Youngbluth (1976) suggests that, although strongly controlled by physical parameters of the environment, some species may vary their vertical distribution and diurnal vertical migration in response to the prevailing food supply. More evidence of the significance of the migration for feeding purposes is given by Youngbluth (1975) who presents his own data on euphausiids and reviews the work of others such as Ponamareva (1971) and Roger (1973b). There are several instances reported of euphausiids closely associated with

5.

V E R T I C U DISTRIBUTION AND MIGRATION

441

chlorophyll maximum layers suggesting that the migration in those cases is a feeding response. Hu (1978) found that the active migrators Thysanopoda monacantha and T . aequalis (including T . astylata) feed a t night whereas the much less actively migrant Nematobrachion sexspinosum feeds during the day. A fourth species Thysanopoda pectinata, which performs a moderate migration, feeds continuously by day and night. Roger (1975), on the other hand, examining the feeding rhythms of tropical species, concludes that the migrations are not for feeding. Youngbluth (1975) argues that euphausiids are probably lethargic at day time depths, especially when living in oxygen deficient layers, and that energy expenditure during the vertical phase of the migration may be less than previously considered (Vlymen, 1970; Klyashtorin and Yarzhombek, 1973). Return from the surface layers to day time depths may be effected primarily through passive sinking (Rudjakov, 1970). Consequently, Youngbluth suggests that residence during the d a y in oxygen deficient layers is probably an adaptation to conserve energy and that McLaren’s (1963) hypothesis is re-enforced. McLaren pointed out that vertically migrating organisms can feed a t night in the higher environmental temperatures of the surface layers. They would then return bo the colder deeper layers during the day where the food would be metabolized and converted into growth. This would, in effect, provide an energy bonus (McLaren, 1963; Mauchline and Fisher, 1969). Rudjakov (1970) questions the statement of many authors that the downward migration of planktonic organisms is performed by active swimming. He reviews the results of measurements of sinking velocities of anaesthetized animals and shows that these velocities are equal to or greater than the velocities observed in actively downward migrating animals. He correlates the presence of body lipids with non-migratory species or components of a population and suggests that evolutionary trends in pelagic organisms are towards reduction of energy expenditure and maintenance of the body in a lethargic state through achievement of neutral buoyancy. Consequently, he argues that a diurnal vertical migration was not evolved by pelagic organisms but is a disadvantageous character of their ancestors. This hypothesis is not supported by the views of Jagersten (1972) on the evolution of the holopelagic life cycle through neoteny. Here, a benthic adult had a pelagic larva that, through evolutionary processes, adopted adult characteristics and dispensed with the benthic adult phase of the life cycle. The pelagic adult, however, undoubtedly

442

THE BIOLOGY OF EUPEIATJSIIDS

had different environmental requirements from those of the larva. It would probably, for instance, require food particles of a larger size. Consequently, the original ontogenetic migration, with an amplitude of the same order of magnitude, would be maintained although shifted upwards in the water column. Such ontogenetic migrations are clearly present in the life cycles of many modern euphausiids. Diurnal vertical migration could conceivably have evolved, Z;hrough selective pressures, as a further development of the onbogenetic migrations. Rudjakov argues that the more specialized euphausiids Stylocheiron species do not perform diurnal vertical migrations and that this fact supports his hypothesis that there are evoluijionary trends dispensing with such a migration. The species in this genus that do not migrate (Table IV) live in the epipelagic zone where their food is plentiful and a marked vertical migration probably not of any advantage to them. Evolutionary trends towards a lethargic life style for purposes of energy conservation are suggested by the work of Childress (1977) in the context of the mesopelagic and bathypelagic environments. The significance of these trends is probably to allow colonization of the deeper environment, with its restrictive nutritional resources, rather than to enable the organisms to dispense with a diurnal vertical migration. A further aspect has been identified by Miller (1970), who suggests that planktonic organisms, like their terrestrial counterparts, can be divided into two broad categories: diurnally active (also nonmigrating) and nocturnally active (also migrating). These adaptations, 'including the evolution of a diurnal vertical migration, allow the pelagic community to exploit, to the full, the three-dimensional environment of the oceans, a conclusion supported by the work of Roger (1975).

CHAPTER 6

FOOD AND FEEDING Mauchline and Fisher (1969) review the feeding methods and food of euphausiids and Nemoto (1 972a) has provided a historical account of such investigations. The feeding appendages have been described in some detail and their role in food gathering studied. Weigmann-Haass (1977) has produced excellent drawings of the mandibles, maxillules and maxillae of the larvae of Euphausia hanseni. She found that the larval mandibles have an asymmetric lacinia mobilis and a pair of processus incisivus accessorius (spine row) that are absent in the mandibles of the adults. These structures are present in the mandibles of the calyptopes of Nematoscelis dificilis (Gopalakrishnan, 1973) and of all larval stages of Euphausia gibboides (Knight, 1975) and E . fallax (Knight, 1978). The lacinia mobilis appears to be absent but the processus incisivus accessorius is present on the mandible of calyptopes of Thysanopoda tricuspidata (Knight, 1973). These -processes, a feature of the mandible of species of bhe Peracarida, do not occur in adulti Eucarida (Mauchline and Fisher, 1969). The above authors also describe the appendages of the larvae in detail. Until recently less attention was paid to the form and function of the mandibles which, unlike other appendages, are noti directly concerned with the filtering of food from the water. Their function is to macerate and grind the food. Preliminary studies of the mandibles of adult, euphausiids suggested that the presence of a large pars molaris, the grinding region, was associated with a filter feeding habit (Mauchline and Fisher, 1969; Weigmann, 1970; Casanova, 1974 ; Ponomareva, 1976a; Nemoto, 1977). The following index, relating the size of the pars molaris to the size of the pars incisiva, cutting region, of the left mandible was used by Nemoto (1977): Index =

Width of pars molaris Length of pars incisiva

The regions measured are shown in Fig. 17. The mandible has a complex shape and consequently there is a variety of measurements, especially of the pars incisiva, that can be made. The mandibles of

Ipm:

'I

H

... .::

. . ..............:;.. ... . .

_____

....... ........ ............ i

'i-*

.

..........-: ..-..-_ ... .-._ . -

-

A

__--

. ...... . . . .. ;.:.+?. -----

H

............ . . . . . . . . :;=' ...... ._ .... ._ -_...-

E

;1:11-

H

.-

..

-38.5.

e-

- -..

i..J

-

K

FIG.17. The left mandible A, of Bentheuphausia amblyops showing the two measurements, pia and pib, made of the pars incisiva, a, total length of pars incisiva; b, measurement of main blades only. The measurement of the pars molaris (pm) is shown on the adjacent lateral view of the mandible. The hatched lines on the other mandibles show the measurements made of the pars incisiva. B, Thysa,nopoda acdtifrons; C , Meganyctiphanes norvegica; D, Euphausia krohni; E, E . superba; F, Thysano&sa rmchi; G , T . longicaudata; H, Nematoscelis megalops; J, Nematobrachion boopis; K, Stylocheiron elongatum; L, S. longicorne; M, S. abbreviatum; N, S. maximum. All bars indicate 0.4 mm except that of S. Zongicorne which shows 0.2 mm.

6.

445

FOOD AND FEEDING

Euphuusia superba were ,examined and measured (Fig. 17) and the index calculated (Table V ) ; its value is the same as that determined by Nemoto. Standard deviations of the mean indices quoted by Nemoto have been calculated (Table V) by abstracting the data from Nemoto's graph of length of pars molaris on length of pars incisiva (Nemoto, 1977, fig. 11). The indices found in Bentheuphausia amblyops and Thysanopoda acutifrons when the entire length of the pars incisiva TABLEv. RATIOSOF LENGTHS OF Pars Molaris TO Pars Incisha MANDIBLEOF VARIOUSSPECIES Species Bentheuphausia amblyops Pm/Pia PmlPib Thysanopoda acutijrons Pm/Pia P4Pib T . tricuspidata T . cristata T . cornuta T . egregia T . spinicaudata Meganyctiphanes norvegica Euphausia krohni E. superba Thysanoessa raschi T . longicaudata Nernatoscelis megalops Nematobrachion boopis Stylocheiron longicorne S. elongatum S. abbreuiatum S. m x i m u m

IN THE

n

Present investigation

n

(1977)

Feeding type

10 10

0.52 +_ 0.04 0.75 f 0.07

8

0.71 f 0.03

0-F 0-F

10 10

0.40 +_ 0.05 0.52 0.07

6 7 2 10 8 1

0.64 f 0.08 0.78 +_ 0.08 0.49 0.51 k0.06 0.34+-0.03 0.41

12

1.09 f 0.08

10 10 3 10 10 10 10 10 10 5 10

+-

0.74 k 0-06 0.70 f 0.07 1.08 f 0.04 0.97 k 0.10 0.85 0.03 0.69 f 0.06 0.29 f 0.05 0.31 _+ 0.04 0.45 0.07 0.27 f 0.02 0.42 k 0..04

+

Nemoto

LEFT

0-F P-0 P P P P-0 F-0 F-0 F-0 0 0-P P P

P P

The number of observations and standard deviations are shown, pi* rtnd pib are the two different measurements of the pars ineiaiva explained in Fig. 17. F, filter feeder; 0, omnivorous feeder; P, predatory feeder.

(Fig. 17, pi") is measured are different from those calculated by Nemoto (Table V). Close agreement is obtained in Bentheuphuusia amblyops by measuring only the length of the main cutting edge (Fig. 17, pib). Meaiurement of this length in Thysanopoda acutij'rons still produces a different index from that found by Nemoto (Table V). A comparison of the mandible of this species, shown in Fig. 17, with that illustrated by Nemoto suggests that mandibles of North Pacific individuals are

446

THE BIOLOGY OF EUPHAUSIIDS

of a slightly different form from those of north-east Atlantic individuals. Kulka and Corey (1978) describe an ontogenetic change in the morphology of the mandibles of Thysanoessa inermis that is correlated with changes in its feeding behaviour and distribution; the pars molaris is relatively smaller in 0-group animals (neglecta form) than in I-group animals (predominantly inermis form). The indices calculated in this investigation and by Nemoto are shown in Table V. The feeding type, determined by analyses of stomach contents (Table VI) is also shown. The first type of feeding is considered dominant where two are shown, e.g. O-F is predominantly omnivorous but also filter feeds to a considerable extent. There does indeed appear to be a correlation between a low index and a predominantly carnivorous habit as Nemoto suggests. Weigmann (1970) figures the mandible of Euphausia diomedeae and, from measurements of her drawing, this mandible has an index of 1.12; this species is considered to be omnivorous (Table VI). Artiges et al. (1978) determined values for the “edge index” of the mandibles, an index developed by Itoh (1970) in studies of the functional morphology of the mandibles of copepods. Two euphausiids, Meganyctiphanes norvegica and Euphausia krohni were used, indices being calculated for tihe mandibles and for the pars incisiva alone. This index, in the case of euphausiids, is less easy to use and interpret than that of Nemoto. Berkes (1975) has re-examined the function of the feeding appendages, although inconclusively. According to him, there axe conflicting views on how the food basket, the space present between the knees of the thoracic legs and mouth parts, functions. The food basket is frequently full of material that has probably entered it, inadvertently to some extent, while the euphausiid has been in the densely packed plankton bucket. Much of this material is probably not food in the normal sense, and the contents of the food basket are consequently treated with some reserve when defining the diet of a species (Casanova, 1974). The term “food basket” is not usually used in the context of filter feeding because there is little evidence that this region functions in such a manner during filter feeding. Prtvlov (1971a, b), however, states that the diatoms and other material filtered from the water by Euphausia superba form a bolus in the food basket. Maceration by tihe mouthparts separates the edible liquid material which is swallowed from the rest of the material which is washed out of the basket by the euphausiid before the next period of filtration. Consequently, Pavlov is suggesting %hatperiods of filtration alternate with periods of sorting and maceration of the food. This has not been reported previously but may be a common mode of filter feeding.

6.

FOOD AND FEEDING

447

The term food basket is Blways used in the context of describing the feeding of euphausiids on sea bottom material and on living prey, such as copepods. In these situations, it functions as a collecting point for material being macerated by the mandibles. Berkes (1975) has re-examined the mechanism of filter feeding in Meganyctiphanes norvegica, Thysanoessa raschi and T . inermis. He attempts to identify the activity of the pleopods and thoracic legs in the generation of the currents but also considers the contribution that might be made by the motion of the animal when it swims. His conclusions appear to differ little from those expressed by Mauchline and Fisher (1969). The feeding and respiratory currents of Euphausia superba are illustrated by Pavlov (1971a) who suggests that, in addition t o the normal feeding currents, the pleopods can produce a current from in front of the animal, drawing water posteriorly through the food basket. Pavlov (1976b) suggests further that the antenna1 s d e s function to direcb the feeding currents. This is the first time that the functional morphology of these appendages has been considered. Similar forms of scales are present in other decapods and, of course, in the Mysidacea. They are flat appendages, usually with naked outer margins but with setose inner margins. They probably do induce directional flow in feeding currents as Pavlov suggests but may also act, in conjunction with the telson and uropods, as “diving planes”. The structure of the thoracic legs is discussed by Mauchline and Fisher (1969). The setulation of the seven pairs of legs of Meganyctiphanes norvegica and the six pairs of Euphausia krohni is described in some detail and figured by Artiges et al. (1978). Those of the former species are more heavily setose although smaller particles are probably retained by the finer sefae of E . Erohni. Species in the genera Thysanoessa (some species), Tessarabrachion, Nematoscelis, Nematobrachion and Stylocheiron have one or two pairs of the thoracic limbs elongated. The function of these limbs is unknown but is usually considered to be associated with carnivorous feeding. Berkes (1975) observed the use of the elongated second pair of limbs of Thysanohsa inermis f. neglecta. They were flexed to press lumps of detrital material against the mouthparts. Subsequently, Berkes (1977b) shows that the neglecta form feeds more extensively on crustaceans while the inermis form consumes more “mush”. Laboratory experiments demonstrated that the neglecta form readily consumed Artemia nauplii while the inermis form did not. Descriptions of the stomach, its form, structure and histology are given in Mauchline and Fisher (1969). Supplementary descriptions of

448

THE BIOLOGY OF EUPHAWSIIDS

the gross shape of the sto.mach of Bentheqhuusia amblyops, Thysanopoda cornuta, T . spinicaudata, T . egregia, T . cristata, T . acutifrons, T . monacantha, T . aequalis and T . tricuspidata are given by Nemoto (1977). The walls of the stomach of the three bathypelagic species, T . cornuta, T . egregia and T . spinicaudata, expand when filled with food but are folded when empty. There is a large variety of spines forming the internal armature of the stomach. The food of some 20 species, in addition to that of the 29 specie8 described by Mauchline and Fisher (1969), has been studied (Table VI). Determinabion of the diet of a euphausiid is difficult. The stomach contents frequently, if not usually, contain unrecognizable amorphous material often termed detritus, a description thab is probably incorrect in many cases. This material is frequently the macerated ‘internal tissues of animals and plants and fragments of setae, chaetae or diatom frustules are mixed with it. Species living in shallow regions often feed on sea bed material and consequently mud or sediment particles then occur among this material (Mauchline and Fisher, 1969; Berkes, 1976). Items in the left half of Table VI-detribus, algae, diatoms, dinoflagellates, tintinnids, Radiolaria, Foraminifera, coccolithophores-are probably obtained predominantly through filter feeding, while ibms listed in the right half are probably obtained through carnivorous methods of feeding. As Berkes (1975) pointed out, carnivorous feeding of euphausiids is probably “encounter-feeding” because, as Mauchline and Fisher (1969) state, there is no evidence of hunting or stalking of prey. This is further confirmed by the experiments of Fowler et al. (1971a). They fed Meganyctiphanes norvegica on Artemia. The euphausiid swam towards Artemia of 3-6 mm length in a twisting and turning manner with the thoracic legs rapidly opening and closing. This opening and closing produces a region of negative pressure around the mouthparts and water in front of the animal is drawn between the legs to the mouthparts. Fowler et al. state that this activity continued until the euphausiid “touched and subsequently trapped the Artemia between the appendages”. The euphausiid frequently repeated this process until three or four Artemia had been trapped and then it slowed its swimming activity and started t o ingest the Artemia. It is often very difficult, on the basis of analyses of stomach contents, to decide whether a species is a filter feeder (F)j an omnivore (0)or a predator (P).No species probably feeds by one method alone. Some, like Euphausia superba, probably filter feed almost exclusively throughout the season when phytoplankton is plentiful (Pavlov, 1971b). Euphausia triacantha was found by Pavlov (1976a) to feed

6.

FOOD AND FEEDINU

449

on E . superba during the winter months when its normal food was scarce. Other species may feed exclusively by one method on a patch of phytoplankton for a period of hours or days then change to an omnivorous diet or to a strictly carnivorous diet for a period. Diurnal changes in diet have been demonstrated, Meganyctiphunes norvegica being often predatory on small copepods a t depth during the day but filter feeding on phytoplankton or dinoflagellates in the surface layers at night (Mauchline and Fisher, 1969). Roger (1973a), like many previous authors (Mauchline and Fisher, 1969), points out that the organisms most likely to be recognized among the stomach contents are those with the most resistant structures, resistant principally to the mechanical maceration of the mouthparts and the armature of the stomach walls. Pavlov (1971a, b) found that Euphausia superba eats the large spiny diatoms belonging t o the genera Chuetoceros and Rhizosolenia. These diatoms are easily destroyed and if.the stomachs are not examined quickly after capturing the euphausiids, then only the more damage-resistant species, such as those in the genera Fragillariopsis, Coscinodiscus etc., are found. Further, some structures, such as the luminescent organs of copepods of the genus Pleuromumma, are easily recognized as belonging to tt specific genus, while other copepods that may occur in the diet may not have distinctive structures by which they can be recognized among the stomach contents. Consequently, a species list of dietary components can be extremely misleading without the addition of quantitative assessments of the importance of individual items. There are many difficulties present in producing such quantitative information, not the least of which is the relative importance of the “detritus”, “green mush” or other amorphous material that occurs so commonly. Such material may represent the internal tissues of copepods that have been extracted by the mouthparts which have then rejected the exoskeletons. There is no obvious way of quantifying, in a meaningful way, the amount of this material relative to the other materials among the stomach contents. Roger (1973s) attempted to identify all material in the stomachs of tropical euphausiids as plant, animal or originating from both. It is even difficult to do this at times. Future investigations may produce methods of quantifying individual items among the stomach contents, possibly through biochemical analyses and identification of biochemical signatures of some components of the diet. The diurnal diet, however, probably consists of more than one meal. Roger (1973b) found that active feeding by tropical species never ceased completely during the 24 hour period. There was, however,

TABLEVI. FOOD

Species Benth,euplmusia amblyopa Thysanopoda monacantha T . cristata T . tricuapidata T . aequalis T . astylata T . obtumfrons T . pectinata T . orientalis T . microphthalma T . acutifrons T . cornuta T . egregia T . spinicaudata Meganyctiphanes norvegica Nyctiphanes couchi N . australis N . capeneis N . simplex Pseudeuphausia latifrons P . sinica Euphauaia americana E. eximia E. krohni E. mutica E. brevis E. diomedeae E. recurva E. superba E. vallentini E. lucens E. frigida E. paciflca E. m n a E. cryst"l1orophia-s E. temra E . similia E . similis var armata E. mucronata E . sibogae E . distinguenda E. lamelligera

I-

Feeding Dino type DetriDia- jlagel- Tin- Radio(see text) tus Algae toms lates tinnids laria

0-F P-0 P-0 0-F

X

X

X

X

X

X

X

X

X

x

X

X

x

X X

P-0

x

P-0 P

+ microplankton

X

X

P P P

P-0 F

X

X

X

X

x

X

X

X

X

X

X

X

X

X

X

X X

F-0

x

F-0

x

X

F-0 0

x

X

X

X

X X

X X

X

X

X X

X

X

X

X

X

X

X

X

X

X

F F

X

F

X

X

X

X

X

X

X

X

0

X

X

X

X

OF

EUPEAUSIIDS

Fora- CoccoChae- Echmini- litho- Medu- tog- ino- Mol- Amphi. Crust- Ommafera phores sae natha derrna luacrr pods acea tidia Fish X X

X

X

X

X

X

X

X

X

X

X

1, 7 1, 7, 12 7 1, 2, 7, 8

X

X

1, 6, 7, 8, 12

X

X

X

X

X

X

X

X

X

X

X

X

X X

X X

X

X

X X

X

X

X

X X

X

X

X

X X

X

References

X

X

7, 12 7, 8 1 1, 11 11 11 1, 6, 9 1, 6 1 1 1, 2, 5

X

6

X

1

X

X X X

X

X

X

X

X

2, 8 1

X

X

X

X

X

1, 5, 8

X

2

contd.

TABLEVI

.Species

E. gibba E. gibboides E. fallax E. sanzoi E. pseudogibba E. paragibba E. hernigibba E. spinifera E. bnaeni E. longhostria E. triumntha Teesarabrachionoculatum Thysano&sa spinifera T . lOngipe8 T . inapinata T . inemis IT. longicaudata T . parva T . gregaria T . vicinu T . mucrura T . raachi Nenaatoscelia &&ilia N . Wg&pS N . tenella N . rnicrope N . atlantica N . lobata N . gracilis Newtobrachion jtexipea N . sexepinosurn N . boopie Stylocheiron carinatum S. afine 8.suhrni S. rnicrophthalw S. insulare S. elongaturn S. indicurn S. longicorne S. abbreviaturn

s. T n a x h m

Feeding Dinotype DetriDia- &elTinti- Radio(see text) tua Algae t o m late8 nnids laria

F X

X

X

X

F-0

x

X

X

X

X

X

X

X

F

X

X

X

X

0

X

X

X

X

F

X

F-0

x

X

0-P 0 0-P P

x

X

P P

X

X

X

X

X

X

X

X

x

X

X

X

X

X

X

X

X

X

X

X

P

P P P

X

X X

X X

X X

8. robuaturn References: 1, Mauchline and Fisher, 1969; 2, Weigmann, 1970; 3, Pavlov, 1971b, 1974; 8, Ponomareva, 1975, 1976b; 9, Berkes, 1976; 10, Pevlov, 1976a; 11, Nemoto, 1977;

-contd. Fora- CoccoChae- Echmini- litho- Med- tog- ino- Mol- Amphi-Cruat- Ommafera phores w a e mathe d e r m luacs pods acea tidia Fish

References 7

X

X

X

X

X

X

8

X

7, 8 6, 8

X

1

X

X

10

X

X

X

1

X

X

X

X

X

1, 9 1, 9 8 6

X

X

X

X

X

X

X

1, 9

X X X

X X

X

X X

X

4, 7, 8

X

7 X

X

X

x

1,12 1, 7 8 8 8

8

X

X X X

4, Ponomsreva, 1971; 5, Enomoto, 1972; 6, Casanova, 1974; 7, Roger, 1974c, 1976; 12, Hu, 1978.

454

THE BIOLOGY O F EUPHA’USIIDS

a sinusoidal rhythm present. I n general, the most active period of feeding was at night between 2000 and 0600 h but another active period was between 1400 and 2000h. The rhythm in most species was not strictly synchronous with the day-night alternation (Fig. 18). For instance, Nematoscelis tenella and Nematobrachion boopis fed most actively between 1200h and 0000-0200 h. He has integrated his data on the periodicity of the most active feeding of nine species t o show the fluctuating intensity relative to time and depth (Fig. 19). The period of most intense feeding by these species in the surface layers between 2000 and 0600 h was attributed to the six diurnally migrating species, Thysanopoda aequalis, T . tricuspidata, T . orientalis,

Tbysonopoda monacanfha Tbysanopdo pectinafo Nemafobrocbion boopis Tbysanopodo crisfofo Nemotoscelis teneNo

............ ......

0 .

. . . .@

0.0..

DQY

800-400

0.

300-0

000-400 300-0 000 - 300 800 - 300 500- 200

Night

T . monacantha, T . pectinata and Euphuzcsia diomedae. The second most intense period of feeding, at about 400-500 m depth between 1200 h and 2000 h, was mainly attributable to Thysanopoda aequaiis, T . pectinata, T . cristata, T . tricuspidata, Nematoscelis tenella and Nematobrachion boopis. Three of these species, namely Thysanopoda aequalis, T . pectinata and T . tricuspidata, were kherefore feeding while a t depth during the day and also during their vertical migration to the surface layers where they continued feeding through the night (Fig. 18). The first six species (Fig. 18) are migrant, the last three non-migrant. Each species hap a period, usually about 10-14 h when it feeds less actively (Fig. 18). It probably, however, fills its stomach more than once during the 10-14 h when it is feeding most actively and, of course, the diet may change from a carnivorous diet at depth to a herbivorous diet at the surface.

Time of day

0800

1600

1200

B

~

.

~-l.o

n ..,....... ......../, .....,... ._..( ... ..............

2000

m 1 0 - 5 0

2400

50-100

0400

100-1000

FIU.19. Integrated feeding activity of the nine species of euphausiids whose diurnal rhythms are shown in Fig. 18. The data are presented as numbers of actively feeding eupheusiids per 1000 m3 water against time of day and depth (after Roger, 1973b).

456

THE BIOLOGY OF EUPHAUSIIDS

Roger (1975), studying 23 species, concluded that, in general, feeding rhythms are a generic characteristic ; Euphausia species feed mostly a t night, Thysanopoda species tend to feed continuously, Xtylocherion species feed during daylight hours and Nematoscelis and Nematobrachion species feed mostly between noon and midnight. Meganyctiphanes norvegica had a weak feeding rhythm dependent upon body size and season (Fig. 20). Young individuals (0-group) tended to feed more actively at night during the winter but in the summer fed most actively between noon and midnight. The older individuals (I-group) tended to feed most actively during the day in winter and during the night in summer. Thysanoessa raschi tended

90

00

1600

2400

0800

Time of day

90

-

.cd

0o

,

-*u’

-0-group animals - - - I -group animals

,,*’

,

1600

2400

0000

Time of day

FIG.20. Percentage o f Meqanyctiphanes norvegica and Thysanoeasa raschi with food present in the stomach at different times of the day during winter (upper graphs) and summer (lower graph) (after Mauchline, 1960, 1966).

to feed more actively between noon and midnight in winter, but the feeding rhythm is very weak. Ponomareva et al. (1971) found that a diurnal feeding rhythm was virtually absent in one year old but that two year old Euphausia diomedeae in the Indian Ocean feed mosB intensively during the night and morning. The Antarctic E . superba has a circadian rhythm of carbohydrate metabolism, being a nocturnal feeder (Mezykowski and Rakusasuszczewski, 1979). Earlier information on diurnal feeding rhythms is given in Mauchline and Fisher (1969).

Roger (1975) found that species with more pronounced feeding rhythms usually did not have very pronounced diurnal vertical migrations whereas actively migrating species had weak or non-existent feeding rhythms. Ponomareva (1971), on the other hand, found that

6.

FOOD AND FEEDINQ

467

Nematoscelis microps and N . gracilis, which were non-migrants in the Indian Ocean, showed no feeding rhythms. The non-migrant Nematobrachion sexspinosum fed during the day, the migrant Thysanopoda monacantha and T . aequalis fed a t night and the moderate migrator, T . pectinata had no feeding rhythm (Hu, 1978). The diets of young Meganyctiphunes norvegica and Thysanoessa raschi are different from the diets of older individuals (Mauchline and Fisher, 1969). Older and larger individuals of both species eat more crustaceans, such as copepods, and chaetognaths than younger individuals. The younger individuals appear to filter feed to a much greater extent. Weigmann (1970), examining the diets of Euphuusiu diomedeae, E. sibogae ( E . distinguenda) and E . tenera, found that crustacean remains were more common in the stomachs of larger individuals. She found that tintinnid species were selected on the basis of size and form but that there was no comparable selection of the other items in the diet. Fowler et al. (1971a) noted that Meganyctiphanes norvegica fed more readily on larger rather than smaller Artemia. Nemoto and Saijo (1968) and Nemoto (1970b, 1972b) have measured total pigments-chlorophyll a and phaeopigments-in the stomach contents of several species of euphausiids. As expected, measurable quantities occurred in the stomachs of epipelagic species but they also occurred in the stomachs of the bathypelagic Bentheuphausia amblyops and the two mesopelagic species, Thysanopoda cristata and T . pectinata. Further, they found them present in the stomachs of predatory species of the genera Nematoscelis and Nematobrachion. The source of these pigments in these euphausiids was not recognized although it may be through feeding on herbivores or on faecal pellets. A migratory species, such as Thysanopoda pectinata, may, of course, feed directly on phytoplankton in the surface 300 m layer a t night. Sameotio ( 1976a) detected concentrations (sound scattering layers) of the species Meganyctiphanes norvegica, Thysanoessa raschi and T . inermis in the Gulf of St Lawrence by using an echo sounder. Simultaneous net samples provided information on numbers of euphausiids per m3 and estimations of concentrations of chlorophyll a within 5 m of the surface were made. Concentrations of chlorophyll a and the presence of euphausiids were significantly correlated. The Thysanoessa species were probably feeding directly on the phytoplankton and the associated small herbivores. Wroblewski (1977) discusses the effect that euphausiids may have in creating patches of phytoplankton. Several euphausiids have been maintained successfully in the laboratory for varying periods. Artemia eggs and nauplii, with or

458

THE BIOLOGY OF EWPHAUSIIDS

without the addition of algal cultures, form the principal diet offered. Ponomareva et al. (1971) fed, Euphausia diomedeae, E . brevis and Thysanopoda tricuspidata the green alga Chlamydomonas, a peridinian Amphidinium klebsii, the diatom Biddulphia catenata and bacteria either aggregated on particles or singly in suspension. Parsons et al. (1967) found that Euphausia pacijica in the laborabory could consume particles as small as 5 pm, a range of algae of differing cell sizes being offered. Meganyctiphanes norvegica was maintained in the laboratory on a diet of juvenile Artemia of 2-4 mm length and a dilute suspension of Phaeodactylum tricornutum (Fowler et al., 1971a). I n rearing Nematoscelis dificilis, Gopalakrishnari (1973) fed them on Artemia nauplii of about 0.4 mm length and mixed phytoplankton consisting of Lauderia. borealis, Coscinodiscus granii, Cyclotella nana and Coccotithus huxleyi. Larvae of Meganyctiphanes norvegicus and Nyctiphanes couchi were reared on a diet of Artemia nauplii and Phaeodactylum tricornutum (Le ROUX,1973a, 1974).

CHAPTER 7

CHEMICAL COMPOSITION Much more information about the chemical composition of euphausiids is available in addition to that given in Mauchline and Fisher (1969). This is primarily a result of an increased interest in the commercial exploitation of populations of euphausiids as a source of food for man and domestic animals. Grantham (1977) has reviewed in detail the chemical composition of the potentially most important commercial species, the Antarctic Euphausia superba. The bodies of euphausiids can be divided into several major biochemical factors (Table VII). The data in Mauchline and Fisher (1969) have been partially integrated with these additional data. The estimation of water content is subject to considerable experimental error (Raymont et al., 1969, 1971a) and this has contributed in part to a reported range of water contient of 70-80% wet weight. Consequently, some authors, for example Raymont et al. (1969, 1971a), Ferguson and Raymont (1974), Bamsbedt (1976) and Roschke (1978), have expressed these biochemical fractions in terms of percentage dry weight rather than wet weight. This can result in wrong conclusions being drawn from the data (Childress, 1977); lipid, protein and ash are the major constituents of the dry weight and a decrease in one causes an apparent increase in the other two. Childress cites the conclusions of Raymont et al. (1971a) who found, on a dry weight basis, that increased lipid content in Meganyctiphanes norvegica in the autumn was apparently associated with a decreased protein content. Childress recalculated their data on a wet weight basis and showed that in fact the protein content did not decrease in the autumn but, like the lipid content, was at a seasonal maximum. Bamstedt (1976) has produced a seasonal study of the changes in the major biochemical fractions of the dry weight of M . norvegica but it is difficult to assess the significance of all his results without the corresponding information on fluctuating water content. Raymont et al. (1971a, b) found evidence of seasonal fluctuations of water content of some 5 % in M . norvegica and 7% in Euphausia superba, while Harding (1977) shows that larger T h y s u n o h m raschi contain

TABLEVII. QUANTITIES Species Meganycti.phane.3 norvegica Eupha& mperba E . paci&a E . krohni Teasarabrachion oculatunz Thysanobaa racrchi T . inemia

Water 81 +_ 5 73-85 82.8 f 3.8 79.7-80 78.7 81.1-87.5 78.5+5

OF THE

Ash 3.3-4-3 2.3-3.7 14-3.1 3.7-3.8 1-73 1.94-5.60 3.87

MAYORBrocmancluc, CONSTITUENTS IN Lipid

Carbohydrate

Protein

3.8-4.9 1.1-7.0 04-2.4

0.44 0.84

64-12.5 9.9-17.7 10.5-16.7

1.28

0.09

5.9-14.0 8.93

4.88

yo WET WEIGHT Chitin

0.92 0.23-2-45 0.22-0.28

0.5-0.8

Reference, 5 , 8, 9, 14, 20 4,5,7, 11,12,13, 16-19 2, 5, 6, 7, 15 1, 5 3, 6 3, 10, 16, 20 20 ~~~~~~~~~

References: 1, Nicholls et aZ., 1959; 2, Suyama et al., 1966; 3, Fujika et al.. 1969; 4, I(ryuchkOV8and Maksrov, 1969; 5, Mauchline and Fisher, 1969; 6, Omori, 1969; 7, Pierce et al., 1969; 8, Raymont et al., 19698, 19718; 9, Idler and Wiseman, 1971; 10, Ikeda, 1970, 1972; 11, Sidhu et al., 1970; 12, Reymont et al.. 1971b; 13, Yanme, 1971,1974a; 14, Tyler, 1973; 16, Childress and Nygeard, 1974; 16, Yanese, 1975; 17, Roschke, 1976, 1978; 18, Watanabe et d..1976; 19, Granthem, 1977; 20, Hopkins et al., 1978.

7. CHEMIOAL COMPOSITION

461

relatively more water than smaller individuals. Body length, however, is used by Bamstedt as a substitute for wet weight and he was able to demonstrate seasonal fluctuations in the dry weight and lipid content of two theoretical Meganyctiphanes norvegica of carapace length 6 mm and 9 mm. He also examined the chemical compositions of juveniles, males and females separately and detected small differences. The five biochemical fractions were examined on a dry weight basis in Thysanoessa inermis by Raymont et a2. (1969) who found values similar to those in Meganyctiphanes novegica. The chemical composition of the cast integuments of Euphausia pmijka and Thysanoessa species are referred to in Mauchline and Fisher (1969, pp. 270-271); values for dry weight, carbon, nitrogen and organic content are given. Clarke (1976) found that cast integuments of Euphausia 8Uperba represented 3% of the body dry weight, considerably lower than the previously reported value of 6 1 0 % in Euphausia pacifica and Thysano&sa species. Sameoto (1976b) found that the casts represented 6.2f2*2Y0 of body dry weight in Meganyctiphanes norvegica, 5-7 & 1.7% of body dry weight in Thysanoessa inermis and 5.0 & l.Oyo of body dry weight in T . raschi. The dry integuments of Euphausiu superba consisted of 49% ash, 4.2 i-1.4% lipid, 0-045 & 0.02y0 soluble carbohydrate, 0.065 & 0.03% insoluble carbohydrate, and 46.7% protein plus chitin (Clarke, 1976). Yanase (1975), working on the same species, made a more detailed investigation of the casts, examining the chitin, calcium, magnesium and phosphorus contentu; he estimated the casts to be about 35% of the wet weight. Estimations of carbon and nitrogen have been made on several species (Table VIII) and carbon/nitrogen ratios have been calculated, using mean values where applicable. Jawed (1969) determined the relationship between the particulatie nitrogen content (N) and the body dry weight ( W) in Euphausia pwiifica, based on 18 animals whose range in dry weight was approximately 15-19 mg, as:

N

= 0.122

w - 0.090

Carbon, nitrogen and carbon : nitrogen ratios were measured in Nematoscelis megalops from slope water and from Gulf Stream cold core rings by Boyd et al. (1978). Carbon ranged from 15% to 50% body dry weight and nitrogen from 3.5% to 16% body dry weight. The nitrogen and phosphorus contents of Meganyctiphanes norvegica averaged 9.5% and 0.8% of body weight respectively (Roger, 197813). Few elementary analyses of euphausiids have been made and the data of Nicholls et ul. (1959) on Euphuusia krohni were inadvertently

462

THE BIOLOGY O F EUPHAUSIIDS

TABLEVIII. CARBONAND NITROGEN, IN yo DRYWEIGHT,AND CARBON/ NITROUEN RATIOSIN EUPHAUSIIDS Species

Carbon

39.8 1 Thysanopoda aequalis fifeganyctiphanes 33.40-37'00 norvegica

Nitrogen 10.51 9.90-10'80

CIN Ratio

References

3.8

Nemoto et al. (1969)

3.4

39.38 f 0.49 46.50 & 0-33 44-99 & 2-60 35.80

11.47 f 0.22 9.63 f 0.36 8.45 f 1.62 6.80

3.4 4.8 5.3 5.3

41.73 39.45

3.7 3.7

38.10-45.10

11.20 10.63 11.66 10.76 10.40-13.00

38.70 40.07

10.70 12.27

3.6 3.3

E . nana 41.95 E . airnilis 43.15 TLssarabrachion 47.20 oculatum

10.44 10.99 10.00

4.0 3.9 4.7

Mauchline and Fisher (1969) Mauchline and Fisher (1969) Mayzaud (1973a) Mayzaud (1973a) Hopkins et al. (1978) Mauchline and Fisher (1969) Nemoto et al. (1969) Nemoto et al. (1969) Jawed (1969) Yanase (1974a) Mauchline and Fisher (1969) Omori (1969). Childress and Nygaard (1974) Nemoto et al. (1969) Nemoto et al. (1969) Omori (1969)

5.20-7.10

winter spring Euphawia krohni E . mutica E . recurva E . superba E . pacifica

Thysanobsa inermis T . raschi Nematoscelis dificilis

N . atlantica N . microps N . megalops

3.6

42-63 43.92 f 2.81

9.43 9.59 f 1.75

4.5 4.6

Nemoto et aZ. (1969) Hopkins et al. (1978)

40.64 4.38 42.00-43 '70

10.20 & 2.11 10.60-1 1.10

4.0 3.9

Hopkins et al. (1978) Hopkins (1968)

41.76 40.68 42.89 43.66 32.5

10.20 10.69 11-34 10.59 10.7

4.1 3.8 2.6 2.4 3.0

Nemoto et al. (1969) Nemoto et al. (1972) Nemoto et al. (1972) Nemoto et al. (1972) Boyd et al. (1978)

omitted by Mauchline and Fisher (1969). Further new information has been integrated with the earlier data in Table IX. Modern chemical analyses of organisms have to take account of pollution of the environment. Much pollution is primarily restricted to coastal regions but elements introduced through fallout processes, for example, radioactive isotopes from weapons testing, or substances

7. CHEMICAL COMPOSITION

463

TABLE IX. ELEMENTARY COMPOSITION OF EUPHAUSIIDS IN mg/100g DRYWEIGHT

Element

H B Na Mg

Al

P K C& Ti

Euphausia superba

Euphausia pacifica

8.2 1253-1829 94

1690

1780-2430 1076-1400 782-749

2050 1000 762

+ +

+

940

Other species

7600" 14*2b

345 6.1 1160 770-1960

0-17 0.84

Cr

Fe

~~

6600-7360

6521-7300

v

co Ni cu Zn Se Sr Mo Cd Pb

Euphausia Meganyctiphanes krohni norvegica

+ + 12.9 + +

12.8

+ +

8.4-12-4

0.09 0.11 11.2

0.12-1 * 7 14-15 0.36-0.7 1 0.9- 1 8 1-9-11*5 5'3-23.4 0.36 1,1-5

-

6*5*

0.32 0.02-0.66 0.05-3-7

0.055

0.37 ~

10.5- 19.3'

O*lld

~~

Data from: Nicholls et al., 1959; Fujika et al., 1969; Mauchline and Fisher, 1969; Omori, 1969; Halcrow et al., 1973; Leatherland et al., 1973; Mayzaud, 1973a; Small et al., 1973; Yamamoto et al., 1973; Childress and Nygaard, 1974; Belloni et al., 197615; Fowler and Benayoun, 1976; Cutshall et al., 1977. Teasarabrachion omlaturn. b E~phcczl~ 8irnili8. &~ Thy8ano98a raachi end Tes8arabrachion ocuhtum. d Mixed species. + Elements detected qualitatively b y K v c h i and Kano (1975).

such as polychlorinated biphenols (PCB) and organochlorine pesticide residues (DDT residues) are more widespread. Concentrations of elements such as cadmium, arsenic, chromium, zinc, mercury, lead and others in marine organisms contain, in many cases, a contribution from industrial wastes. The degree of pollution of different sea,areas varies in time and space and so variable quantities of individual elements and substances have been recorded in the bodies of euphausiids. Experimental determinations of the rates of uptake and degree of accumulation of various pollutants by euphausiids have been made. The most useful way of presenting these data is by classifying them under the various individual elements and substances.

464

THE BIOLOGY O F EUPHAUSTIDS

Chromium. Watanabe et al. (1976) measured 0.02 mg chromium/ 1OOg wet weight Euphausia superba from 56"33'S, 20'08'E in January, 1973. Manganese. The radioactive isotope 54Mnwas accumulated slowly by Meganyctiphanes norvegica directly from the water, a significant portion being adsorbed on the surfaces of the integument. It was eliminated quickly from the body tissues and cast integuments when the animals were maintained in 54Mn-freewater; a small portion, 1% of the original body burden, was retained after 65 days (AntoniniKane et al., 1972; Fowler et al., 1972). Stable manganese was estimated as 0.2 mg/lOO g dry weight Thysanoessa raschi and as 1.4 mg/100 g dry weight Tessarabrachion oculatum by Fujika et al. (1969). Halcrow et al. (1973) determined the manganese content of Meganyctiphanes norvegica in the Firth of Clyde where extensive deposition of ferromanganese oxides takes place. The euphausiids contained 4-2-141 mg manganese/lOOg dry weight, values much higher than those of 0.20.95 mg manganese/lOO g dry weight found in the same species in the Mediterranean by Belloni et al. (197633). Iron. Accumulation of the radioactive isotope 59Fe directly from the water was slow and significant quantities were adsorbed on the surfaces of the integument of Meganyctiphanes norvegica (AntoniniKane et al., 1972; Fowler et al., 1972). Accumulation of 59Fe from labelled food was variable, probably because of the varying physicalchemical state of the iron. Loss of adsorbed 59Fe from cast integuments placed in 59Fe-free water was slow. Jennings (1977) has studied the distribution of 55Fe, originating from fallout from weapons testing, in the eastern Pacific Ocean by using euphausiids as indicator organisms. Cobalt. Accumulation of the radioactive isotope 5 7 Cby ~ Meganyctiphunes norvegica can take place directly from the water or by way of food. Cobalt is adsorbed from the water onto the surfaces of the integument (Antonini-Kane et al., 1972; Fowler et al., 1972). The release of 57C0from cast integuments placed in 57Co-freewatier was slow. Kuenzler ( 1969b) found that Thysanopoda tricuspidata eliminated cobalt accumulated in the body at only l%/h. Copper. Reeve et al. (1976, 1977) found that Euphausia pacijka is extremely sensitive to concentrations of 5 pg copper/l sea water in controlled environment pollution experiments. Zinc. Detailed studies of the accumulation, retention and elimination of zinc by euphausiids have been made by Fowler et al. (1969, 1971b, 1972), Small (1969), Antonini-Kane et al. (1972), Small and Fowler (1973) and Small et al. (1973). Appreciable quantities of zinc

7. CHEMICAL COMPOSITION

465

are accumulated a t a relatively fast rate directly from the water and at a slower rate from the food. It is adsorbed from the water on the surfaces of the integument, this portion being eliminated from the body burden of the animal a t moulting. Cross et al. (1968) and Fowler et al. (1970) maintained Euphausia p c i $ c a and Thysanoiksa spinifera in sea water containing sbZn and in ebZn-freesea water but on a diet of 65Zn labelled Artemia. They showed, by autoradiographic techniques, that assimilated 65Zn was localized in the integument and muscles; it also occurred in the eyes. Most of the assimilated zinc was located intercellularly and not intracellularly. Its biological half-life in the tissues of E u ~ h u u s ipacijica ~ was relatively long, 140 days (Fowler et al., 1971b). The rate of elimination was exponential (Small et al., 1973), initial rates in s5Zn-free water being high as Kuenzler (1969a) suggested. The main route of elimination depends upon the original source of the isotope to the animal. Zinc accumulated directly from the water is lost a t moulting, the major part of the body burden being adsorbed on the integument, and some is also exchanged with the 65Zn-freewater. Zinc accumulated from labelled food is primarily eliminated from the body in the faecal pellets, although some is lost at moulting and by exchange processes with the 65Zn-free water. Euphausiids readily accumulate zinc and Antezana and Fowler (1972) used Euphuusia mucronata as an indicator organism in a study of the 65Zn Contamination of the south-east Pacific in the immediate period of the 1966 French nuclear weapons test. Arsenic. A concentration of 0.01 mg arsenic/100 g dry weight Euphausia superba was recorded by Watanabe et al. (1976)at 56'33'5, 20'08'E in January, 1973. Kennedy (1976) found 0-18k 0.04 mg arsenic/lOO g wet weight (approximately 0.90 mg/100 g dry weight) in euphausiids off northern Newfoundland. The highest concentration recorded is 4.2 mg arsenic/100 g dry weight Meganyctiphnes norvegica from the entrance to the Mediterranean (Leatherland et al., 1973). Selenium. A detailed study of the accumulation, retention and eliminatjion of selenium by Meganyctiphanes norvegica was made by Fowler and Benayoun (1976). It was more readily accumulated from food, than directly from the water, and its turnover time in the tissues was relatively rapid, its biological half-life being estimated as 37 days. Strontium. Tolkach and Gromov (1975) found that Euphausia superba concentrated by a factor of 4-13 over the concentration in an equal weight of water. Yttrium. 25% of the OOY was adsorbed irreversibly on the surface of the integument of Euphausia superba, the remaining 75% being accumulated in the internal tissues (Tolkach and Gromov, 1976).

466

THE BIOLOGY OF EUPHAUSIIDS

Zirconium-Niobium.. The fission products 05Zr/05Nbwere detected in Euphausia superba in the Scotia Sea in 1967 and 1968 by Tkachenko and Patin (1971) and were probably obtained from radioactive fallout material from weapons testing. Ruthenium. This element forms stable, soluble complexes when mixed with sea water. Its accumulation by Meganyctiphanes norvegica depends on its physical-chemical state (Antonini-Kane et al., 1972; Keakeij et al., 1972). Ruthenium from sea water is absorbed on the surfaces of the integument. The source of ruthenium to the marine environment is radioactive wastes disposed to the sea and fallout material from weapons testing. Tkachenko and Patin (1971) identified ruthenium-106 and rhodium-106 in Euphausip; superba collected in the Scotia Sea in 1967 and 1968. Cadmium. Benayoun et al. (1974) studied the uptake, retention and elimination of cadmium in Meganyctiphanes norvegica. It is accumulated directly from the water or from the food. Assimilated cadmium is located primarily in the viscera but also occurs in the eyes, muscle and integument. Natural cadmium occurred ab concentrations of 0.011 mg/lOO g wet weight Euphausia superba (Watanabe et al., 1976), 0.07 mg/lOO g dry weight Meganyctiphunes norvegica (Benayoun et al., 1974) and 0.023 mg/100 g wet weight of mixed species of euphausiids (Cutshall et al., 1977). Antimony. A concentration of 0.0037 mg/lOO g dry weight occurred in Meganyctiphunes norvegica taken a t the entrance to the Mediterranean in July, 1970 (Leatherland et al., 1973). Caesium. The biological half-life of caesium assimilated by Euphuusia paci$ca was estimated as 6 days by Fowler et al. (1971b). Radioactive 13'Cs, originating from fallout from weapons testing, was detected in Euphausia superba in the Scotia Sea in 1967 and 1968 (Tkachenko and Patin, 1971). Cerium. Studies of the accumulation, retention and loss of cerium by euphausiids have been made by Antonini-Kane et al. (1972) and Fowler et al. (1971b, 1973). The biological half-life of assimilated cerium is estimated as 7.5 h. Cerium is adsorbed from the water on the surfaces of the integument and subsequently lost a t moulting. Mercury. Watanabe et al. (1976) measured 0.001 mg mercury/100 g wet weight Euphausia superba at 56'33'5, 20'08'E and Nagakura et al. (1974) found 0.0008-0.0015 mg/lOO g wet weight in the sam0 species at 59'40'S, 23'05'W, both observations in January, 1973. Knauer and Martin (1972), examining euphausiids off the Californian Coast a t 36'46'N, 122'01'W, found 0.0005-0.0021 mg mercury/100 g wet weight, while Kikuchi and Kano (1975) measured 0.004 mg

7. CHEMICfi COMPOSITION

467

mercury/100 g wet weight in Euphausia pacifica from an undefined norvegica, at the entrance to the Mediterlocation. Megany~ti~hanes ranean, contained 0.026 mg mercury/100 g dry weight in 1970 (Leatherland et al., 1973). Lead. Watanabe et al. (1976) found 0.58 mg lead/100 g wet weight Euphausia superba a t 56'33'5, 20'08'E in January, 1973. Halcrow et al. (1973) found a range of 1.9-3.7 mg lead/100 g dry weight Meganyctiphanes norvegica in the Firth of Clyde where a source of pollution exists. The radioactive isotope 210Pbis a decay product of uranium and occurs naturally in the marine environment; Holtzman (1969) measured 3.75 f 0.33 pCi 210Pb/100g wet weight E . pacifica and 1.32 ? 0.02 pCi 210Pb/100g wet weight Meganyctiphanes norvegica. Concentration of 2loPb takes place in the hepatopancreas (Heyraud and Cherry, 1979). Polonium. The radioactive isotope 210Po occurs naturally in the marine environment. It is accumulated by euphausiids and was found to concentrate in the digestive tract and hepatopancreas (Heyraud et al., 1976a, b; Heyraud and Cherry, 1979), but was not adsorbed significantly from the water on the surfaces of the integument. Folsom and Beasley (1973) found 640 dpm/kg wet weight E . pacijica as 2loPo while Holtzman (1979) measured 26.7 ? 1.3 pCi 210Po/100g wet weight E. pacijica and 5.29 ~f:0.34 pCi 21OP0/100 g wet weight Meganyctiphanes nwvegica. Half the accumulated 210Po was lost from the hepatopancreas of M . norvegica in 6.6 f 0-9 days (Heyraud and Cherry, 1979). Plutonium. Plutonium enters the marine environment as a pollutant from the nuclear power and weapons industry. Fowler and Heyrrtud (1976) suggest that surface adsorption directly from the water on to the integument is the principal method of accumulation of 23'Pu. No information is available about assimilation of plutonium in the internal tissues of euphausiids, but Higgo et al. (1977) measured 0.007-0-010 pCi 2s9+240Pu/100 g wet weight in whole Meganyctiphanes norvagica in contrast to concentrations of 0.03 pCi/ 100 g integument and a range of 0-9-7.0 pCi/lOO g faecal pellets. Uranium. Miyaje et al. (1970) found 1.7-0.2 x mg/100 g dry weight of euphausiids as uranium. Natural levels of alpha radioactivity. These were measured in mixed species of euphausiids and in Nematoscelis megalops and Thysano&sa gregaria by Shannon (1972). He found a range of 9-60 pCi alpha activity/100 g wet weight euphausiids. D D T . Concentrations of DDT residues present in Meganyctiphanes norvegicn, Thysano&sa raschi and T . inermis in the Gulf of S t Lawrence

468

THE BIOLOGY OF EUPHAUSIIDS

during the period June, 1973t o May, 1974 were determined by Sameoto et al. (1976). A comparative study of the concentrations of DDT residues and lipids suggested that the euphausiids metabolize the lipids without decreasing the body burden of DDT residues. Higher concentrations of DDT residues were accumulated by Meganyctiphanes norvegica than by the Thysanoika species. The DDT residues were assimilated from food at efficiencies comparable to those of carbon assimilation (Cox, 1971). Larger Euphamia pacifia retained DDT residues longer than smaller individuals; Cox examined the possible effects of moulting, surface volume ratios and varying diet on the uptake of DDT residues. PCB. Takagi et at?. (1976) found about 0.008 mg PCB/lOO g wet weight E. pacijica in the north-west Pacific; this concentration was higher than that occurring in the predators of E . ppaciJica, namely salmon and trout. Meganyctiphanes norvegica had 0.004-26.0 mg PCB/ 1OOg dry weight during the period November, 1974 t o March 1976 a t a position 5 km off Villefranche-sur-Mer, France (Elder and Fowler, 1977). The faecal pellets contained higher concentrations, ranging from 0.48-3.8mg PCB/lOOg dry weight; this represents, on a wet weight basis, a concentration factor of 1-6x 108 over the concentration in the surrounding sea water. Calorific values of Meganyctiphanes norvegica, Euphausia krohni, E . superba and E . pacijica are given in Mauchline and Fisher (1969). They range from 7Cb122 kcal/100 g we%weight. Tyler (1973) found a mean annual calorific value for Meganyctiphanes norvegica of 94-96 and an overall range of 84-104 kca1/100 g wet weight; these are comparable to the range found by Bratbelid and Matthews (1978) in this species in Western Norway. Hopkins et al. (1978)noted average calorific values of 78.8 kcal in Meganyctiphanes norvegica, 65-2kcal in Thyaanoasa rmchi and 92.7 kcal in T. inermis, all per 100 g wet weight. The seasonal change in the calorific value of Meganyctiphanes norvegiw is illustrated by Brattelid and Matthews (1978) who found a Winter maximum of about 110 kcal and a spring minimum of about 96 kcd/ 1OOg wet weigh$. The calorific value of Eqhausia superba was 100110 kca1/100 g wet weight (Chekunova and Rynkova, 1974). Childrms and Nygaard (1974) calculated the calorific value as 106 kca1/100 g wet weight in E. pacijica. The lipids of euphausiids have been investigated in some detail in recent years. Mauchline and Fisher (1969) present data on the lipid content of 26 species; additional information is given in Table X. H6rring (1972) estimated a range of 1.6-6.9% wet weight as lipid in 17 north-east Atlantic species. He found 2.4% wet weight as lipid in

TABLEX. LIPIDS(yo WET WEIGHT)IN SPECIESOF EUPHAUSKIDS ~

SpeCier,

Thyaanopoda mnacuntha T . tricwpidata T . aequalis T . obtwifrons T . microghthalm Meganyctiphanea norvegica

E . superba

E. paci&x

E . eimilia E. gibboidas Thysanohaa longipea T . inermis

T . raschi Nemtoacelk micropa Nematobrachion b o w N . sexapinoaum Stylockiron abbreviatum

Month

20°N, 23OW 0ct.-Nov.

% lipid

Reference

1.1

Morris (1971)

Nov. Oct.-Nov. Oct. Nov. Apr.June

1.1 1.2 1.4 1-3 1.8-3.6

Morris (1971) Morris (1971) Morris (1971) Morris (1971) Sameoto et al. (1975)

0ct.-Nov.

1-6-2.4 1.9-2.1 1.0

Ackman et al. (1970) Morris (1972a, b, 1973) Morris (1971)

Australia 20"N, 25"W Oct.

1.8 0.8

Bishop et al. (1976) Morris (1971)

-

-

27ON, 15"W 27"N, 15"W 41"N, 2OW S. Tasman Sea Ross Sea 66"S, 70"W Enderby Land 42"N, 146"E

Jan.-March Feb. July May

0.9-1-1 11.7 2.1 1.1 6.1

Morris (1972a, 1973) Culkin and Morris (1969) Morris (1971) Morris (1971) Sidhu et al. (1970)

April Dec.-Jan. Dec.-Feb.

8.0 3.0 3.4

Sidhu et al. (1970) Raymont et al. (1971b) Tsuyuki and Itoh (1976)

Sept.

1.4

Japan

March

1.3

Takahashi and Yamada (1976) Takahashi and Yamada (1976) Kayama et al. (1976) Childress and Nygaard (1974) Kayama et al. (1976) Morris (1971) Morris (1971) Kayama et al. (1976)

17"N, 27"W 20°N, 23"W 33"N, 14OW 19"N. 26OW E. Canada E. Canada

Nyctirphanee wuchi N . auatralis Euphwia amerimw E. lcrohni E . brevia

~~

Sea area

-

-

4loN, 2"W

July

Japan Sea May, Oct. S. California

1.1-3.6 3.1

Japan 22"N, 23"W 28"N, 14"W Japan Sea

2.1 1.2 1.2 7.7

June Oct. Dec. May, Oct.

G. of St. MayJune Lawrence N.W. Oct. Atlantic Japan May

1.0-10-2 Sameoto et al. (1975) 1.8

Ackman et al. (1970)

5.7

G. of St. May-June Lawrence 17"N, 25OW Nov.

14-4.0

Takahashi and Yamada (1976) Sameoto et al. (1975)

2.4

Morris (1971)

lSON,25"W Nov.

2.7

Morris (1971)

30"N, 23"W April 25"N, 17"W Oct.-Nov.

1.0 1.7

Morris and Sargent (1973) Morris (1971)

470

TEE BIOLOGY OF EUPRAUSIIDS

Thysanopoda cristata. The other 16 species investigated have also been examined by other workers whose results are closely similar and quoted by Mauchline and Fisher (1969) or shown in Table X. There was no correlation between lipid content and modal day depth of occurrence of the species. There are seasonal variations in the quantities of lipids stored by epipelagic species in middle and high latitudes, maximal concentrations occurring in the autumn and winter, minimal in the spring or summer after egg production ; this results in a seasonal change in calorific value, noted above. Larger Meganyctiphanes norvegica store relatively more lipid than smaller M . noruegica (Mauchline and Fisher, 1969) and this has been confirmed by Raymont et al. (1971b); no such relationship between body size and lipid content was found in Thysunoessa raschi (Mauchline and Fisher, 1969) or in Euphausia superba (Raymont et al., 1971b). The average lipid content of an euphausiid is in the range 0.5-2-5% wet weight. Thysanoessa species have frequently been found with %lo% wet weight as lipids (Table X ; Mauchline and Fisher, 1969). Other species that have been recorded with a lipid content in this range are Thysanopoda aequulis, Euphausia krohni, E . brevis, E . superba and Stylocheiron maximum. Some other species may also have a high lipid content and may not have been analysed yet or sampled at the appropriate season. Seasonal and other aspects of lipid storage are discussed in Mauchline and Fisher (1969). The lipids can be divided into several fractions such as triglycerides, &glycerides, monoglycerides, steryl esters, wax esters, sterols, free fatty acids and phospholipids. Some of these fractions have been quantified in the lipids of Meganyctiphunes norvegica, Nyctiphanes australis, Euphausia superba, E . pacijica, E . similis, Thysanoksa inermis, T . raschi, T . longipes and Nematobrachion sexspinosum by Ackman et al. (1970), Van der Veen et al. (1971), Morris and Sargent (1973), Lee (1974), Bottino (1975), Sargent and Lee (1975), Bishop et aZ. (1976), Kayama et al. (1976), Takahashi and Yamada (1976), Tsuyuki and Itoh (1976) and Watanabe et al. (1976). The dominant fractions are the phospholipids, free fatty acids and triglycerides. Wax esters assume importance in some species but not in others (Sargent, 1976). Bottino (1973, 1975) found 20-44% of the lipids as wax esters in Euphausia crystallorophias but they were not pxesent in E . superba. Takahashi and Yamada (1976) record 30% of the lipids of ThysanoGsa inermis as wax esters but only found traces in Euphada pacijica. Morris (1942a) found 5% of the lipids to be wax esters in Meganyctiphanes norvegica and Ezlphawi krohni compared with estimates of 15-30% in Thysanopoda aequalis and T . microphthalma.

7. CHEMICAL COMPOSITION

47 1

Morris and Sargent (1973) determined 35% of the lipids of Nematobrachion sexspinosum t o be wax esters, while Kayama et al, (1976) record 1-3-5% wax esters in Euphuusia pacijica and E . similis and 7.7% in the lipids of Thysanoessa longipes. Sargent and Lee (1975) found 3% of the lipids of T . raschi to be wax esters. Tropical samples of Thysanopoda tricuspidata, T . obtusifrons, Euphausia recurva and Nematobrachion sexspinosum contained 7-21 % of the lipids as triglycerides but had only traces of wax esters present (Lee and Hirota, 1973). Several comparative studies of the occurrence of wax esters in pelagic organisms have been made and are referred to by Bottino (1975) and Kayama et al. (1976). These studies have suggested that deeper living animals store energy as wax esters rather than as triglycerides and that this form of storage aids in attaining neutral buoyancy and so a lethargic life style. Most information on the occurrence of wax esters in euphausiids can be used 60 support this hypothesis, except in bhe case of Euphuusia crystallorophias. This species has wax esters in preference to triglycerides and is not considered to live in deep water; it lives in high latitudes in the Antarctic, probably, for the most part under the ice. Bottino (1975) suggests that the monoenoic waxes of this species may have an adaptive role in this extreme environment ; Rakusa-Suszczewski and McWhinnie (1 976) measured the freezing point of the body fluid as - 2.13"C in the course of a study on the resistance of Antarctic fauna to freezing. Analyses of the fatty acid and alcohol compositions of the wax esters of E. crystallorophias, E. pacijica, Thysanoessa inermis, T . longipes and Nematobrachion sexspinosurn are given by Morris and Sargent; (1973, Bottino (1975), Kayama et al. (1976) and Takahashi and Yamada (1976).

Analyaes of %hefatty acids present in the lipids of Meganyctiphunes norvegica, Euphuusia superba, E . pacijica and ThysanoEssa raschi are given in Mauchline and Fisher (1969). Many further analyses have been made, both of the fatty acids and alcohols, in a range of species: Thysanopoda monacantha, T . tricuspidata, T . uequalis, T. obtusifrons and T . microphthulma by Morris (197 1) ; Meganyctiphanes norvegica by Ackman et al. (1970), Morris (1972b, 1973) and Belloni et al. (1976a); Nyctiphanes couchi by Morris (1971); N . awtralis by Bishop et al. (1976); Euphausia americana by Morris (1971); E . krohni by Morris (1973) ; Euphausia americana by Morris (1971) ; E . superba by Hansen (1969), Pierce et al. (1969), Hansen and Meiklen (1970), Sidhu et al. (1970), Van der Veen et al. (1971), Bottino (1972, 1973, 1974, 1975)) Tsuyuki and Itoh (1976) and Watanabe et al. (1976); E . pacijica by Pierce et al. (1969), Van der Veen et al. (1971)) Lee (1974), Kayama

472

THE BIOLOGY OF EUPHAUSIIDS

et al. (1976) and Takahashi and Yamada (1976); E . crystallorophias by Bottino (1974, 1975); E . gibboides by Morris (1971); Thysanoessa longipes by Kayama et al. (1976); T . raschi by Sargent and Lee (1975); Nematoscelis microps by Morris (1971) ; Nematobrachion boopis by Morris (1971); N . sexspinosum by Morris and Sargent (1973); Stylocheiron abbreviatum by Morris (1971). All these analyses are similar in gross characteristics to those shown in Mauchline and Fisher (1969). Small variations in the detailed fatty acid profiles exist between species and within species. Morris (1971) found evidence of a seasonal variation in the fatty acid composition of Euphausia gibboides and that deep living non-migrant species may have higher proportions of monounsaturated acids. Other variations reflect the fatty acid composition of the diet, probably more so in the case of triglycerides than the complex lipids (Bottino, 1974). Investigations into the fatty acid composition of the phospholipids have been made (Ackman et al., 1970; Morris and Sargent, 1973; Bottino, 1975; Takahashi and Yamada, 1976; Tsuyuki and Itoh, 1976). The major components of the phospholipids are phosphatidyl choline and phosphatidyl ethanolamine. Earlier information is given in Mauchline and Fisher (1969). Cholesterol, reported as the major sterol in Euphausia pacijica and representing 0.03% wet weight in E . superba (Mauchline and Fisher, 1969) was present a t a concentration of 0.063% wet weight in E . superba according to Watanabe et al. (1976). The cholesterol content of Meganyctiphanes norvegica was 0.09% wet weight (Idler and Wiseman, 1971). Cholesterol and provitamin D were estimated in the sterol fraction of Euphausia superba by Vinogradov and Kandiuk (1967). Amino acid composition of the proteins of several species of euphausiids have been determined primarily t o show that they contain a balanced group of essential amino acids. The species investigated are shown in Tables X I to XIII. Watanabe et al. (1976) compared the amino acid content of raw and cooked Euphausia superba and demonstrated a loss of free amino acids in the cooked samples. Mauchline and Fisher (1969) discuss, in considerable detail, the occurrence of vitamin A, its isomers and the carotenoid astaxanthin in several species. Watanabe et al. (1976) found 2-2-2.7 pg vitamin A/g and 1.6-7.8 pg astaxanthinlg wet weight Euphausia superba; these values are within the ranges given by Mauchline and Fisher (1969). The concentration of tocopherol (vihmin E) in E . superba was l6-78pg/g wet weight (Watanabe et al., 1976). Yanase (1971) found the following quantities of vitamins, expressed .as pg/g wet

TAELEXI.

-0

ACID COMPOSITION OF PROTEIN HYDROLYSATES OF EUPHAWSIIDS

Suyama et al. (1965)

Kayama et al. (1976)

Euphausia Euphausia Euphausia Thysmoessa pacifica vallentini pacifica longipes (gamin0 acid/lOOgprotein) (g amino acid116 g N or 100 g protein) Cysteic wid Taurine Aspartic acid Threonine Serine

Glutamic acid Proline Glycine Nmine

Valine Cystine Methionine Isoleucine Leucine Tyrosine Phenylalanine Lysine Histidine Tryptophan Arginine Ornithine TOTAL,

mg/g dry weight

yo dry weight

-

-

13.7 4.8 4.8 14.7 3-5 5.4 5.6 5.2 1.4 3-3 5.2 7.8 4.2 6.5 7.8 2.2 1-6 6.0

12.5 4.8 4.7 14.6 3.3 4.9 5.9 5.4 1.2 3.2 5.3 8-4 4.1 6.6 8.6 2.1 1.7 5.9

114-1 1.7

-

4.9-5.0 34-44. 14-9-15.2 3.3-3-6 5.6-5.7 5-9-6.9 4.5-4.9 0.8-1-2 1'9-3.1 4.5-4.6 7-9 4'1-4.5 7.1-7.4 7.1-7'2 2.1-2'6

11.0 4-9 4-5 14.1 3.5 5.5 5.7 4.9 0.6 2.2 5.1 8.2 4.9 8.4 7.1 2-5

5.1-6-4

6.1

Sriniwasagam et itl. (1971) Meganyctiphanes norvegica ( % of -1) -

11.5 4.5 4.1 18.0 3-1 4-1 5.5 5.2

1.1 2.4 4.8 7.5 4.3 4.7 9.6 3.0 -

6.8 0.1 410480 41-48

TABLE=I.

AMINO ACIDS OF PROTEIN HYDROLYSATES OF Euphausiu superba ~

Amino acids

Sidhu et al. S u y a m et al. (1965) (1970) (after Moiseew) Srinivamgarn et al. (1971) Burkholder (g amino acid/ et el. (1967) 16 g N or 100 g ( g amino acid/ (yoamino N ) of protein) 100 g protein) (yoof total) (yoof total)

Cysteic acid Taurine Aspartic acid Threonine S0rine

Glutamic acid Proline Glycine Alanirle Valine Cystine Methionine Isoleucine Leucine Tyrosine Phenylalanine Lysine Histidine Tryptophan Arginine Ornithine TOTAL mg/g

dry weight

% dry weight

a Possibly including isoleucine.

-

1.8

-

-

-

10.4 4.9 4.6 15.7 5-6 8.6 6.9 6.1 1.6 3.5 4.5 8.7 5.2 6-1 9.2 2.4 1.9 7.5 -

13.3 5-0 4.8 18.3 3.8 5.3 6.1 5.3 1.5 2.8 5.7 8.3 5-9 5.1 11.5 2.5 1.3 6.8 -

12.2 4.7 5.0 14.6 4.2 4.7 5.5 5.9 1.5 3.0 5.1 7.8 4.1 6.5 8.6 2.3 1.5 6.2 -

11

10.7 4.3 3.8 15.2 3.3 4.6 5.6 5.2 1.2 2.8 5-1 7-7 4.3 5.0 10.0 3.5 7.5 0.4

7 2 12 6 8 4 15" 7 6 12 2 8

-

423-460 42-46

Ferguson and Raymont (1974) yo of total (9 sampks) 0.5-2.4 0.2-1.4 94-11.5 4.2-5.1 4.5-5.4 11-3-13.4 2.6-3.6 4.2-4.8 5.6-7.0 5.3-5.8 0.1-1.1 24-3.0 5.2-6.2 7.5-8.4 4.2-5.3 4.6-5.2 9.0-10-7 2-5-2.8 6.6-7.7

04-0.8

7.

475

CHEMICAL COMPOSITION

TABLEXIII. FREEAMINO ACIDSOF EUPHAUSIIDS Euphausia superba A k n o acids

Meganyctiphanes norvegica

Srinivasagam et al., (1971) (Yo of t o w

Cysteic acid Taurine Aspartic acid Threonine Serine Glutamic acid Proline Glycine Alanine Valine Cystine Methionine Isoleucine Leucine Tyrosine Phenylalanine Lysine Histidine Tryptophan Arginine Ornithine TOTAL

mg/g dry weight Yo dry weight

1.8 15.9

0.9 3.8 2.1 1.0 8.3" 14.0 10.2 4-1

-

0.5

3.7 10.6 7.7 5.7

4.5 0.8 4.5 8.9 3.7 1.6

0.4"

1.9 2.0" 8.1 7.5

0.8 1.2 1.6 3.3 1.2 11.0 3.3 18.3 7.3

103- 107

20-28

3.1 2.6 3.1 0.9

8.4

10.3-10.7

2.0-2.8

" Not an average value but value for one animal. weight E . superba : riboflavin, 1-58; vitamin B,, 1.1 ; Ca-pantothenate, 0.15; niacin, 0.70; folic acid, 0.66; biotin, 0.10; vitamin BIZ,0.16. The occurrence of two carotenoid pigments, astaxanthin and @carotene, are discussed in considerable detail by Mauchline and Fisher (1969) and little further information is available. Herring (1972) found no positive correlation between increasing body content of carotenoid pigments and increasing depth of occurrence in the 17 species of euphausiids that he examined. The concentrations of total carotenoid pigments ranged from 0 to 173 pg/g wet weight but most values were within the broad range 20-120pg/g wet weight euphausiids. There was a close relationship between body size and pigment content of

476

THE BIOLOGY OF EUPHAUSIIDS

the various species with the exception of Stylocheiron abbreviatum, S. elongatum and S. maximum which had significantly less relative to their body size than occurred in the other species. Herring links this with the reduction of the number of photophores in these species. Astaxanthin alone has an average seasonal range of concentration of 30-80 yg/g wet weight in the two species Meganyctiphanes norvegica and Thysanoessa raschi (Mauchline and Fisher, 1969). A recent more detailed analysis of the carotenoid pigments of Euphausia superba by Czeczuga and Klyszejko (1970) is shown in Table XIV. This analysis TABLEXIV. CAROTENOIDPIGMENTS IN DIFFERENT REGIONS OF THE BODYOF Euphauaia superba AS yo WETWEIUHT(Czeczuga and Klyszejko, 1977) Carotenoid

Astaxanthin Astaxanthin ester Cryptoxanthin Flavoxanthin Zeaxmthin Hydroxy - {-carotene Dihydroxy-[-carotene Unknown TOTAL CONTENT

(pg/g wet weight)

Entire body

Cephalothorax

Abdominal

7.4 17-7 27.8 10.9 13.0 21.7 1.5 -

33.5 9.7 32.8

40.3 14.3 36.6

6-1 11.8 6.1

8.8

15.710

7-855

muack.9

16.124

was made on a single sample collected new South Georgia and suggests that further investigation may demonstrate seasonal changes in the concentrations present. Adenosine triphosphate (ATP), ribonucleic acid (RNA) and desoxyribonucleic acid (DNA) concentrations and their seasonal variations have been measured in Meganyctiphanes norvegica by Skjoldal and Bamstedt (1976, 1977). ATP and RNA were maximal in the spring with a secondary maximum in the autumn in the Korsfjorden, western Norway. DNA was maximal in February, the time when the gonads were actively developing. Homarine is a betaine that is presumed to have an important osmoregulatory function in marine animals. It is N-methyl picolinic acid betaine and was first isolated in lobster muscle. Hirano (1975) measured 2.54 mg homarinelg wet weight in the whole Euphausia superba. The source of homarine is probably the algal diet. The commercial exploitation of euphausiids, especially E . superba and E . pacijica as food for man and animals has necessitated several

7.

CHEMICBL COMPOSITION

477

chemical investigations. Amino acid analyses, mentioned above, have shown that they conform with the nutritional standards of animal feeding stuffs. The treatment of fresh krill immediately they are caught must be rapid because they deteriorate quickly through rapid autolysis (Suzuki and Kanna, 1977); many individuals are damaged by the nets (Kelle, 1978). Three proteases, two of which are trypsinlike and a third of unknown specificity, were extracted from E . superba by Seki et al. (1977) and considered responsible for this autolysis. Five proteases, called A,, A,, B, C and D, were extracted from E . superba by Chen et al. (1978) and their activities and stability studied. Sakasai et al. (1978) have developed a simple method of estimating protease activity in krill extracts. Studies of the effects of freezing and subsequent thawing on the nutritional quality of fresh and cooked E . superba have been made by Kryuchkova and Makarov (1969), Yanase (1974a), Seki et al. (1975) and Suzuki and Kanna (1977). Kelly et al. (1978) found that bacterial counts on fresh and frozen E . superba were low (650-llOO/g a t 25°C) but that *hese bacteria have a high proteolytic and lipolytic activity. The chemical composition of the exoskeleton was examined by Yanase (1975). It can be removed by machine during processing and Yanase (1974b) has described the preparation of edible condensed solubles from E . superba. These solubles have the characteristic flavour and taste of crustacean meat and can be used for seasoning food. The overall nutritional value of krill meat is comparable to that of casein and shrimp meat (Arai et al., 1976).

CHAPTER 8

VISION AND BIOLUMINESCENCE The structure of the compound eyes of euphausiids is described in Mauchline and Fisher (1969) and new information is available. Elofsson and Dahl (1970) examine the optic ganglia of Thysanopoda tricwpidata, Meganyctiphanes norvegica, Euphausia hemigibba, E. gibboides, Nematobrachion Jlexipes and S . abbreviatum and compare their structure with those of other crustaceans. The nuclei of the neurons of the lamina ganglionaris are located to the side of the neuropile instead of in front of it, as in other crustaceans. They state that euphausiid eyes retain, to some extent, an embryological condition, but that the presence of the photophore in the eye stalk may have had some influence in producing this situation. There is no marked difference between the structure of the ganglia of divided and undivided eyes. Nemoto (1969) examines the relative sizes of the eyes of the north Pacific species Bentheuphausia amblyops, Thysanopoda wrnuta, T . monacantha, T . tricuspidata, Euphausia diomedeae, Tessarabrachion oculatum, Thysanokksa gregaria, Nematoscelis difficilis, Nemutobrachion boopis and Stylocheiron affine. He expresses the height of the eye as a percentage of carapace length. Comparable measurements of the eyes of Thysanopoda egregia and T . spinicaudata are given by Nemoto (1977). Measurements of the eyes of species occurring off the west coast of Scotland have been made and combined with Nemoto’s data in Fig. 21. Eyes of the 42 species in the genexa Meganyctiphanes, Nyctiphunes, Pseudeuphausia, Euphausia and Thysanoessa ( T .inermis, T . spinifera and T . raschi only) have heights ranging from about 16% to 30% of carapace length. These species are all epipelagic or coastal (Table IV, p. 431). The eyes of the 13 Thysanopoda species are also circular with height ranging from 4% t o 20% of carapace length and eight of these species are mesopelagic or bathypelagic (Table IV). The remaining 29 species in the genera Thysanokksa (with the excepCion of bhe three species mentioned above), Tessarabrachion, Nemutoscelis, Nematobrachion and Stylocheiron have divided eyes with heights ranging from about 30% to 65% of carapace length. Their vertical

8.

100

VISION AND BIOLUMINESCENCE

l-

I

I

479

I

1

200

40C

E

fa 0

tl

2000 loo0j

T

c

3000

P5

4000

k

s

1

40E

% 20

0

c,

FIG. 21. Relative sizes and shapes of eyes of species living in different ranges of depth. The height of the eyes is expressed aa a percentage of carapace length (scale to lower right of diagram). North-east Atlantic species have been added to the north PacSc species described in Nemoto (1969, 1977).

distribu~ionand behaviour vary between species and genera (Table IV). The size of the eyes relative to body length may vary within a species. Casanova (1977) found that the eyes of Meganyctiphunes norvegica in the Mediterranean have diameters ranging from 1.35 mm

480

THE BIOLOGY OF EUPHAUSIIDS

to 1.90 mm compared with a range of 1.32-1-78 mm in individuals of the same range in carapace length in the eastern Atlantic. The number of ommatidia present in a row tended to be higher in the eyes of Mediterranean than in Atlantic individuals; the numbers of ommatidia in a row ranged from 72 to 90 in the Mediterranean and from 69 to 87 in the Atlantic. This species lives a t greater depths in the Mediterranean where the penetration of sunlight is greater than in the waters of the eastern Atlantic (Casanova, 1977). Mauchline and Fisher (1969) state that the screening pigments of the eyes have not been identified, with the exception of the carotenoid, astaxanthin. Fisher (1967) confirmed the presence of astaxanthin and also melanin and ommochromes in the eyes of Meganyctiphanes norvegica. The structure and function of the photophores in the different genera and species are described in Mauchline and Fisher (1969). The photophores of 33 species have been examined by Herring and Locket (1978). The diameter of the photophores increases with increasing body size of the animals except in the two bathypelagic species Thysanopoda egregia and T . cornuta. Other species in the genus Thysanopoda have markedly smaller photophores relative to body size than species in all other genera; many of the species in this genus are mesopelagic or bathypelagic in habit (Table IV, p. 431). Sexual dimorphism in the distribution and sizes of photophores is discussed earlier (see Table I, p. 378). A study of the morphology and histology of the photophores of adult Meganyctiphanes norvegica, E. gibboides, Nematoscelis microps, Stylocheiron abbreviatum and a species of Thysanopoda was made by Petersson (1968) while Herring and Locket (1978) examine bhose of Euphausia gibboides and Bassot (1966) those of Meganyctiphanes norvegica. Petersson also describes the development of the photophores in furciliae I-V of Meganyctiphanes norvegica. Herring and Locket confirmed, through immersion of live larvae in 5-hydroxytryptamine, that the photophores in the eyes are the first to become functional, followed by that in the f i s t abdominal segment. The first abdominal photophore is able to luminesce even when it is not yet a t a strongly pigmented stage of development. The histological descriptions of Petersson and Herring and Locket are detailed, illustrated with photographs, and confirm, for the most part, the results of the earlier studies. The posterior cell mass, referred to as light-producing cells in Mauchline and Fisher (1969), does not consist of only one type of cell. There are four distinct types in the photophores of M . norvegica and Euphausia gibboides and three in the photophores of the other species. The rods or striations of the

8. VISION AND BIOLUMINESUENUE

481

striated zone appear to be produced by cubical cells of the posterior cell mass, either by secretion or possibly by villi formation. This striated zone or rod mass, however, is a large capillary bed according to Harvey (1977) and is termed a lantern by Herring and Locket (1978). The blood supply to the photophore is most rich in the posterior regions where a nerve net exists. The rods or striations of the striated zone are in three groups; they vary in length, are lanceolate and lie close against each other. Lastly, Petersson observed three distinctly separate layers within the lens of the photophores of Meganyctiphanes norvegica; the lens was previously considered uniform in structure. Harvey (1977) describes it as composed of particles of about 150 nm in diameter with a structure similar t o that of p particles of glycogen. Herring and Locket (1978) also conclude that the lens of Euphausia gibboides is rich in glycogen; they state that the inner mass of the lens appears to be homogeneous. The histology of the pigment layer or reflector and the lamellar ring is also described by these authors. Petersson (1968) and Herring and Locket (1978) examined the mode of nervous control of the luminescence in the context of short emissions (flashes) of light and conclude that the nervous system controls sphincter muscles of the blood capillaries in the posterior cell mass. Constriction of the muscles stops the blood supply, and so the supply of oxygen to the secreted luciferin. A steady flow of blood would result in a luminous glow while a rapid increase in the rate of flow of the blood would produce a sudden flash. Luminescence was inhibited in E . superba when environmental oxygen decreased below 90% saturation (Kils, 1979). Luciferase extracted from Euphausia pacijica and E . sirnilis, as well as from two ostracods and several teleost fish, reacted with the luciferin extracted from tihe toadfish, Porichthys notatus, according to Tsuji et al. (1971). Extracts from Cypridina species and what Yamaguchi (1 975) calls Euphausia japonica cross-reacted but extractis from Oplophorus species and Euphausia japonicu did not react. The spectral characteristics of the luminescence of E . pacijca and T . raschi are illustrated in Mauchline and Fisher (1969). Similar spectral curves have been found by Swift et al. (1977) for Euphausia tenera with a maximal emission at 468 nm and by Herring and Locket (1978) for Nernatoscelis megalops with a maximum emission at 463 k 4 nm and a half intensity band width of 54 nm. Herring and Locket further found that the emission spectra recorded from homogenates of Thysunopoda monacantha, Nyctiphanes couchi, Euphausia brevis, E. arnericana, E . gibboides and E . hernigibba are closely similar, having maximum

482

THE BIOLOGY OF EUPHAlJSIIDS

emission between 467 nm and 473 nm and half intensity band widths of 44-53nm. Mauchline and Fisher (1969)refer to unpublished work of Tett; this has now been published (Tett, 1969, 1972; Tett and Kelly, 1973). A photoflash was used to stimulate the luminescence of Meganyctiphanes norvegica and a delay between the photoflash and the luminescent response quoted as 0.96 s in Mauchline and Fisher (1969, p. 247); this delay should read 96 s. Tett; '(1969)found a comparable delay of 99 & 19-9s in Thysanoessa raschi. This delay time was inversely related 60 temperature. The stimulatory effect of 5-hydroxytryptamine and lysergic acid diethylamide, the inhibitory effect of y amino butyric acid and 21-(3-dimethylaminopropylthio)-cinnamanilide,Squibb 10643 and the non-effect of adrenalin, acetylcholine chloride, L-glutamic acid and sodium salt on the production of luminescence are discussed by Mauchline and Fisher (1969).The effect of 5-hydroxytryptamine (5-HT), its analogues and several other pharmacological compounds on the production of luminescence by Euphausia brevis, E . americana, E . gibboides, E . hemigibba and Nematoscelis megalops has been investig&ed by Herring and Locket (1978). They confirm that 5-HT is stimulatory and also found stimulation by bufotenine (n, n-dimethyl 5-HT), and to some degree, by 5-methoxytryptamine. Adrenergic compounds investigated had no stimulatory effect with the exception of carbamylcholine. The angular distribution of the light emitted from a photophore of Thysanopoda cristata, Megunyctiphanes norvegica, Nematobrachion sexspinosum and Nematoscelis tenella was examined by Herring and Locket (1978).The light is transmitted as beams with an angular distribution closely matching that of the ambient daylight in the sea. Changes in the angular distribution of the light relative to the body axis of the animal is effected in he anterior-posterior plane by rotation of the photophores. Meganyctiphanes norvegica were moved from conditions of light to dark and a seasonally changing proportion of the experimental animals found to respond to the change by luminescing. The majority of the animals respond during the breeding season but nob at other times of the year (Mauchline and Fisher, 1969). Tett (1972)repeated these experiments with Thysanoessa raschi and obtained a similar broad pattern of response. I n addition, however, he demonstrated sexual differences. The proporbion of males responding was greater than that of females in 17 of the 27 months tested. Tett concludes that spontaneous luminescence in T . rawhi is related to breeding behaviour and in parbicular to the transference of spermatophores by

8. VISION AND BIOLUMINESCENCE

483

the males to the females. Gopalalqishnan (1975) discusses the sexual dimorphism of the photophores in Nematoscelis species. He points out that the occurrence of chitinous plates dorsal to enlarged photophores in adult males are specific in pattern and associated with sexual maturity. He suggests that they have a role in species recognition at mating. Ivanov (1969) found brilliantly luminescing swarms of Ewphausia superba. This appears to be the only record of a surface swarm of this species luminescing spontaneously and Ivanov suggests that swarms can thus be located visually at night and an estimation of the commercial value of a sea area made. Raymond and De Vries (1976) observed E . crystallorophias, collected at the sea surface in McMurdo Sound, luminescing in the laboratory but do not state that it was also luminescing spontaneously in the sea. The subtropical Nyctiphanes capensis occurred in an immense aggregation or surface swarm, estimated to have an area of 86.1 km2, a t the sea surface during the night off south-western Africa. This aggregation was luminescing so that “observers aboard the vessel noted a continuous diffuse luminescence (milkiness) whose source was unobservable from the deck” (Cram and Schiilein, 1974) ; phytoplankton was also probably luminescing.

CHAPTER 9

INTERNAL ANATOMY AND PHYSIOLOGY Mauchline and Fisher (1969) describe the internal anatomy and histology of euphausiids. New descriptions of the structure and histology of photophores are referred to in Chapter 8 and descriptions of integumental sensilla found on the carapace are given in Fig. 2 (p. 381). Further descriptions of the distribution of such sensilla and gland openings over the entire integumentis of Meganyctiphunes norvegica and Stylocheiron longicorne are given by Mauchline (1977d) and Mauchline and Nemot,o (1977). The presence of sensilla and gland openings in the integuments of euphausiids are most easily detected by digesting the whole or parts of the animal in hot 10% aqueous potassium hydroxide. The clean integument is then washed in distilled water, transferred t o 70% ethanol and subsequently stained in a 1% solution of chlorazol black E in 70% ethanol. Excess stain is washed off in 70% ethanol and the specimen transferred to distilled water. It is then mounted in polyvinyl lactophenol on a microscope slide. The potassium hydroxide digests the soft tissues of the gland opening or sensillum and leaves a pore or hole in the integument. This pore represents the route of a direct connection between a sub-integumental structure and the outside of the integument. This connection may be (1) the duct of a gland, (2) a tube connecting a sensillum with its axons and sensory neurons located in the hypodermis or (3) a tube through which nerve fibres from the peripheral nervous systiem pass into that part of the sensillum external to the integument. The distributions of such pores in the treated integument have been described but no information is yet available on the structure and function of the sensilla and glands at these sites. Various aspects of the physiology of euphausiids are described by Mauchline and Fisher (1969) and much new information is now available. The earlier work was predominantly concerned with respiration, feeding and moulting. The blood vascular system is described in detail by Mauchline and Fisher (1969). The work of Henderson (1927) was overlooked; she showed that the he& beat of Thysanobsa species

9.

INTERNAL ANATOMY AND PHYSIOLOGY

485

ranged from 125 to 380 beatslmin over the temperature range - 2°C18°C. A recent study of'the ultrastructure of the membrane systems of the cardiac muscle cells of Meganyctiphanes norvegica has shown that it is more similar to that of the cardiac muscles of the other decapods than that of other malacostracan crustaceans (Myklebust, 1977).

Few species of euphausiids live in coastal environments, the majority inhabiting fully saline oceanic water. Sheard (1965) defined the distributions of some tropical species in terms of temperature and chlorinity of the environment. A detailed experimental study of the tolerance of Euphausia pacijca to changes in environmental temperature and salinity has been made by Gilfillan (1972a, b). He used respiration rate as a reference and tested the reactions of individuals from different latitudes, between 30" and 55"N,and at different seasons of the year to changes in temperature and/or salinity. There was no correlalJion between latitude and the capacity of the animal to withstand change; a small seasonal effect was present. There is an interaction between the effects of changes in temperature and salinity, especially of reduced salinity in the presence of elevated temperatures. Gilfillan (1972b) also studied the capacity of oceanic, oceaniccoastal, and coastal groups of E . pacijca to withstand these changes. A gradient of capacity was found, coastal individuals withstanding change much better than oceanic individuals. The effects of hydrostatic pressure on euphausiids has been investigated experimentally. MacDonald et a,?. (1972) subjected Euphausia krohni to increments of 50 atm at 5 min intervals to a maximum environmental pressure of 200 atm. Activity at 1 atm was mainly confined to beating of the pleopods. The animals became more active at 50-100 atm but at 100 atm and 150 atm a violent flexing of the body in the dorso-ventral plane occurred. This flexing resembled the escape reaction except that it was jerky and poorly co-ordinated. Pressures of 200 atm reduced the activity of the pleopods over a period of 1 h. George and Marum (1974) found that in Meganyctiphanes norvegica pressures of 74.8 f 12.5 atm increased the level of activity of the animals, pressures of 229.2 f 13.5 atm induced paralysis and pressures of 387 f 36.5 atm inactivated them. The LD,, was found to be at 262 atm. Furciliae of M . norvegica responded to a decrease in pressure by moving downwards away from light; increasing the light intensity simultaneously with an increase in pressure masked the response t o pressure (Rice, 1967). It is difficult t o interpret the significance of such experiments. The results are frequently not directly comparable because different methods of treatment are used.

486

THE BIOLOUY OF EUPHAUSTIDS

Euphausiids, that is those species that have been experimented with, are considered to be "low resistant species" t o increased pressures (Menzies and Selvakumaran, 1974a, b ; Selvakumaran et al., 1974). Gilfillan (1972a, 1976) finds evidence that i n t r k i c properties of different sea waters of the same temperature and salinity affect the capacity of E . pacijka to withstand stress. He interprets these intrinsic properties as different concentrations of trace metals and organic compounds. The effects of changing concentration of such substances in the marine environment on the horizontal and vertical distributions of planktonic organisms are unexplored. Euphausiids consume 0.04-1-0 pl oxygenlmg wet weightjh depending upon the species and environmental temperature (Mauchline and Fisher, 1969). Most values fall in the range 0-1-0-3 pl O,/mg wet weight/h. Pearcy et al. (1969) found no evidence of a die1 rhythm of oxygen consumption in Euphausia pacijtca. They showed that the weight specific oxygen consumption decreased with increasing body dry weight as did Mayzaud (1973a) for Meganyctiphanes norvegica. The average rate of oxygen consumption was 1 . 3 4 ~ 1O,/mg dry weight/h a t 10°C; this is equivalent to a consumption of about 0.27 p1 O,/mg wet weight/h. Childress (1975) measured a range of 0.08-0.58 p1 O,/mg wet weight/h at 10°C in E . paci@u. Nival et al. (1973) found a range of 1.21-5-5 pl OJmg dry weightlh in E u p ~ krohni, ~ ~ a equivalent to a range of 0.24-1.10 pl/mg wet weight. Mayzaud (1973a) found that Meganyctiphanes norvegica consumed 1.27 f 0.22 p1 O,/mg wet weight/h a t 13°C or approximately 0-25pl 0, on a dry weight basis. A range of 0.5-1.0 pl O,/mg dry weight/h was consumed by M . norvegica maintained at temperatures ranging from 5°C t o 20°C (Mayzaud, 1973b). Sameoto (1976b) determined the following rates of oxygen consumption over the designated ranges of temperature : M . norvegica, 1-35-2-08pl O,/mg dry weight/h over 2"-1OoC; Thysanohsa inermis, 0.62-1.70 pl O,/mg dry weight/h over Oo-15"C; T . raschi, 1.29-1.90 p1 O,/mg dry weight/h over 0"-4"C. Kils (1979) found that Euphausia superba consumed 0.5-1-4 pl O,/mg dry weight/h a t 1°C. The logarithmic relationship between oxygen consumption and body wet or dry weight has been investigated. Herding (1977) found that respiration (R)in pg O,/h was related to wet weight ( W ) in mg by the following equations in Thysanohsa raschi:

R R

=

0-67 WoS2at 5.6"C

= 0.53

WO'82at 1.2"C

9.

INTERNAL ANATOMY AND PHYSIOLOGY

487

He reviews the literature, discusses the physiological significance of tihese equations, and suggests that the weight exponent is constant, at 0.82 in the case of T . raschi. Environmental factors such as temperabure influence the proportionality coefficient only, but he quotes examples where both the weight exponent and proportionality coefficient have been reported to be influenced. He concludes, however, that metabolism is related to some power of wet weight near 0.75 although this value probably is low in the case of euphausiids. Nikolaeva and Ponomareva (1973) determined the following equations for five speciea of the tropical Pacific Ocean and seven species of the tropical Indian Ocean. Respiration (R)is measured in pl 0 2 / h and ( W ) is wet weight:

R R

Pacific Ocean Indian Ocean

=

0.09

= 0.3

We66 at

20°C

WoS3at 20°C

The weight exponent of the Indian Ocean species is near that of Harding-0-82-but that of the Pacific species is much lower. According to Harding (1977), the weight exponent on a dry weight basis is close to 1.0. Chekunova and Rynkova (1974) found the following relationship for respiration rate and body dry weight in Euphausia superba :

R

=

0.121

W 0 ' 9 ' 3 5

Harding examined the relationship between respiration, in pl 0 2 / h and the surface area (AS') of the body, in mm2, in Thysanoiissa rmchi :

R R

=

0.02 SPo0 at 5.6"C

=

0.03 S1'O1a t 1~2°C

Observations of the respiration rate of Euphausia paci$ica in water where the percentage oxygen saturation ranged, in 10% steps, from 10% to 100% were made by Ikeda (1977a). He found that the respiration rate increased with increasing oxygen saturation up to a maximum value in water with an oxygen saturation of 80%. The respiration rate was not affected by levels of oxygen saturation above 80%. Levels of oxygen saturation less than 85-90%, however, severely affect the bioluminescence and rate of respiration of E . superba. Kils (1979) shows that below 85% O2 saturation the respiration rate decreases and no compensation is possible. This species is consequently restricted to well oxygenated layers. Determination of respiration rates of marine animals by incubation for a measured period of time is a relatively lengthy procedure,

488

TIIE BIOLOGY OF EUPHAUSIIDS

subject to experimental errors. Methods of using specific enzyme reactions as rate hdices are being investigated. The mitochondria1 and microsomal electron transport system (ETS) may provide an indirect method of measuring respiration that is both more accurate and more convenient than the present methods (King and Packard, 1975a, b ; Owens and King, 1975). The exploratory investigations, some of which have been made using euphausiids, have shown a close correspondence between ETS activity and oxygen consumption. The only reference to excretion of nitrogen in euphausiids made by Mauchline and Fisher (1969) is to the work of Conover and Corner who found that Meganycltiphanes norvegica excreted 0.19-0.60 pg nitrogenlmg dry weightlday at 3'4°C. More recently Mayzaud (1973~3, b) found that this species excreted 9-52 0.41 pg N/mg dry weightlday in winter but this rate of excretion was only 4.85 1.16 pg N in the spring. About 85% of the nitrogen was excreted as ammonia nitrogen. Jawed (1969)found that Euphausia pacijica excreted, on average, 1.44 and 2.47 pg nitrogen/mg dry weightlday a t temperatures of 4 O C and 10°C respectively. Ammonia was the dominant excretory product (80-85%), followed by amino acid N (11-12-5%) and traces of urea (1-2-1.4%). Jawed (1973) found comparable rates of excretion of nitrogen in E . pacijca and Thysanoessa longipes. Ikeda (1977b) concludes that there is no difference in the rates of excretion of ammonia between starved and fed Euphausia pacijica ; there was, however, considerable variation within his data. Takahashi and Ikeda (1975) measured rates of excretion of ammonia and inorganic phosphorus by E . pacijca while feeding on different concentrations of phytoplankton. They found increased excretion rates with increased phytoplankton density and suggest that the excretion rates of vertically migrating euphausiids would be expected to increase by 1.5-3 times when they enter the phytoplankton concentrations of the euphotic zone. Mayzaud (1973a) discusses the validity and significance of measurements of nitrogen excretion. He found that there were seasonal changes in the rattes of metabolism, chemical composition and the reaction of Meganyctiphanes norvegica to starvation. Seasonal variation in the rates of ammonia and phosphate excretion were observed in M . norvegica by Roger (1978b). The highest rates of 0.25 p g t a t o m s NH,-N/mg/day and 0.026 pg-atoms PO,-P/mg/day occurred in the spring while rates of excretion a t other times of the year approximated to 0.070.11 pg-atoms NH,-N/mg/day and 0.01 pg-atoms PO,-P/mg/day. Lasker (see Mauchline and Fisher, 1969) estimated that Euphausia pacijca assimilated 66-95%, mean 84%, of ingested carbon when fed

9.

INTERNAL ANATOMY AND PHYSIOLOCAY

489

on Artemia. Conover found that Meganyctiphanes norvegica assimilated 65% of the natural organic matter in itis food (misprinted as 6.5% on p. 279 of Mauchline and Fisher, 1969). Fowler et al. (1971a), in feeding Artemia to M . norvegica, found that the percentage of organic matter assimilated from the food was 73.9-91.9%, mean 84.4%, values that are higher than Conover’s. Euphawia diomedeae had an assimilation efficiency of 74% on a diet of bacteria on particles, 82% on a diet of peridinians, and 91% on a diet of green algae (Ponomareva et al., 1971). Moulting of euphausiids is described in Chapter 9 of Mauchline and Fisher (1969). No further work on the physiological aspects of moulting has been done. The data quoted by Mauchline and Fisher (1969), in addition to new data, have been used in studies of growth rates of euphausiids described in the next chapter. Mauchline (1977~) has reviewed present information on the effects of temperature and size on moultring of euphausiids including the recent data of Fowler et al. (1971a, c), Gopalakrishnan (1973) and Le Roux (1973a). The duration of the intermoult period is determined primarily by the body size of the individual and the environmental temperature. At constant temperature, the intermoult period increases logarithmically against body size. Moulting usually occurs at night (Fowler et al., 1971c) but this diurnal rhythm of moulting appears to be weakened at low environmental t,emperatures. Moulting can be shifted from night to day by maintaining the animals in a reversed photoperiod (C. Miller, personal communication of data of Miss B. Dexter). The durabion of the intermoult period decreases with increasing environmental temperature (Fowler et al., 1971c; Clarke, 1976; Sameoto, 1976b). Jennings and Halverson (1971) examined variations in the concentrations of subcuticular acid phosphatase activity during the moulting cycle of Thysanoessa raschi. Highest concentrations are found during the 24 h period within which moulting occurs. They suggest that measurement of this activity in samples of populations would allow assessment of the proportion of individuals moulting or about to moult at the time of sampling.

CHAPTER 10

GROWTH, MATURITY AND MORTALITY Mauchline and Fisher (1969) reviewed the information on growth rates, fecundity and longevity of euphausiids ; detailed analytical studies of ten species were available. New or supplementary investigations of seven of these species and the first study of Nematoscelis megalops have now been made. Studies of the growth rates require measurements to be made of the animals. The various measurements used in the earlier investigations are discussed fully by Mauchline and Fisher (1969) but a variety of measurements, rather than one or two standard measurements, are still used. Several recent biometrical investigations of species are available (Table XV). The regression lines for total body length on carapace length for immature and mature male and female Beganyctiphanes norvegica (see Mauchline, 1960) are compared with those of Matthews (1973) and Hollingshead and Corey (1974) for the same species in Fig. 22. Mauchline measured carapace length as the distiance between the base of the eye and the dorsal median posterior edge of the carapace. The other autihors measured this length as the distance from the base of the eye to the posterior lateral margin of the carapace, a longer measurement. Consequently, their lines are displaced to the right in Fig. 22. Jones (1969) measured total length of Thysanoessa longicaudata as the distance between the tip of the rostrum (instead of the base of the eye) to the end of the telson and does not state how he measured carapace lengbh. Smiles and Pearcy (1971) measured carapace length from the base of the eye to the posterior margin of the carapace, but do not state whethex this margin is lateral or dorsal median. Consequently, it is not possible to compare these lines directly but simply to state that their slopes are closely similar, as expected. There are no accepted standard measurements of euphausiids although total length is usually measured from the base of the eye to the posterior end of the telson, excluding the setae. Roger (1973g, 1978a), like Jones (1969), includes the rostrum in his measurements of total length of euphausiids. Some species have a short rostrum, while others have a long one which is liable to damage. Female

TABLEXV. REGRESSION CONSTANTSOF BIOMETRICAL RELATIONSHIPS Regression constants y=bz+a or log y = b log x a

+

Species Meganyctiphanes norvegica immature J immature 9 mature mature 8

Parameters

T L on C L

a

- 0.74

b

References

c?

0.74 4.09 3.60 3.04

4.44 4.18 3.72 3.61 3.05

Q

8-06

2.22

2.41 0.66 0.29

3-18 2.54 3.06

Mauchline (1960) Mauchline (1960) Mauchline (1960) Mauchline (1960) Hollingshead and Corey (1974) Hollingshead and Corey (1974) Matthews (1973) Smiles and Pearcy (1971) Jones (1969)

2.08 9.33

3.19 1.83

Kulka and Corey (1978) Kulka and Corey (1978)

- 2.30 - 0.05 1.60 1.44 0.90 - 4.31

3.94 3.53 3.21 3.11 4.26 2.58

Mauchline (1960) Mauchline (1960) Mauchline (1960) Mauchline (1960) Fowler et al. (1971a) Mauchline (196713)

- 4.36

2.59

Mauchline (1967b)

- 3.03

3.58

Mackintosh (1973)

- 2.46

3.18 3.10 2.90

Mackintosh (1973) Mauchline (1967b) Nemoto et al. (1977)

3.09 2.85 3.35 3.39 3.34 3.62

Nemoto et al. (1977) Nemoto et al. (1977) Clarke (1976) Mackintosh (1973) Sarhage (1978) Chekunova and Rynkova (1974)

d and ? EuphauAa paci$ca Thysanohsa longicawlata T . inermis 0-group d and 9 I-group 6 and 9 Meganyctiphanes norvegica immature 8 L' on C L immature mature 6 mature 0 L1 on U L 6 log V on log T L 9 Euphausia superba mature Q log V on log T L d and ? EUPHAUSIACEA

Thysanopoda log W , on cornuta log T L Thysanopoda egregia Thysanopoda spinicaudata Euphausia superba

- 5.24 - 4.88

- 5.07 - 4-73 - 2.64 - 2.75 - 2.75 0.51

contd.

TABLEX V - c o n t d . Regression constants y=bx+a or log y -- b log x a

+

Species

Parameters

a

b

References

- 4.52 -5.11 - 1.17

2.58 2.95 2.77

- 3.54

3.60

Mauohline (unpublished) Roger (19736) Matthews and Hestad (1977) Sameoto (197613)

- 0.99

2.92

December

- 0.85

2.91

June

- 0.42

2.35

Euphausia superba

- 1.38

3.81

Thysanoessa inermis Thyaanohsa rmchi Several species together Meganyctiphanes log W, on norvegica log L' log W, on log U L EuphcLzLeia auperba W , on W w

- 3.15 - 3.14 - 2.86

3.38 3.17 2.92 3.31

Brattelid and Matthem (1978) Brattelid and Matthew (1978) Brattelid and Matthew 1978) Chekunova and Rynkovs ( 1974) Sameoto (1976b) Sameoto (1976b) Lindley (1978a) Fowler et a2. (1971e)

- 0.66

3.24

Fowler et al. (1971a)

* 0.09

- 2.27

0.21 0.26

- 0.39

0.87

Clarke (1976) Chekunova and Rynkova (1974) Harding (1977)

0.94

- 2.62

0.12 2.40

Clarke (1976) Harding (1977)

1.11

0.84

Harding (1977)

1.49

0.95

Harding (1977)

EUPHAUSIACEA

Thysano&sa inemis log W , on log C L Meganyctiphanes log W , on norvegica log T L November

Thysanohsa raachi log W, on log w w Euphausia superba W , on W , Thysanoessa raschi log S A on log L log S A on 1% ww log S A on log WIJ

GL,carapace length;

0.13

L1,body length, base of eye to junction of abdomen and telson; S A , surface area of body; TL,total length, base of eye to end of telson; U L , uropcd length; V , body volume; W A ,body ash weight; W,, body dry weight; W , ,body wet weight.

l( -1 1 1 1 1 1 1 5

10

20 Total length (mm)

50

2

6

10

14

Carapace length (mm)

FIG. 22. Left, regression lines relating body wet weight (solid line) and body volume (broken line) to total length of euphausiids. The dots are body weight-length values of species from Roger (1973g), the corresponding regression line being indicated by the dotted line. Right, regression lines relating total length t o carapace length. Meganyctiphane.9 norvegica : 1, short lines are males, long lines females ; broken lines are immature, solid lines are mature (Mauchline, 1960). 2, Males and females of Korsfjorden (Matthews, 1973); 3, males and 4, females of Passamaquoddy Bay (Hollingshead and Corey, 1974). Euphausia pacifica: line 5 (Smiles and Pearcy, 1971). Thysanoema Zongicaudatn: line 6 (Jones, 1969). Regression equations for all lines are given in Table XV.

494

THE BIOLOUY OF EUPHAUSIIDS

Nematoscelis megalops have a long thin rostrum, frequently damaged in sampling, while males of this species have no rostrum. Inclusion of the rostrum in a measurement of body size is inadvisable and makes comparative work difficult. An accurate and complete description of the measurements made should always be given. Roger (1973g, 1978a) determined total body lengtih and the corresponding body wet weight in 20 species of tropical euphausiids. His data are plotted in Fig. 22 and a regression line, whose equation is given in Table XV, fitted. It lies between the regression lines relating body volume and body weight to total length given for Euphausiacea in Mauchline and Fisher (1969), the equations of which are also given in Table XV. As mentioned previously, Roger included the rostrum in his measurement of total length. This is an increase in the normal measurement of total length and would displace his data to the right of the original regression line in Fig. 22. This is true for all except the largest animals, those over 100 mg body weight. This may reflect a tendency for the rostrum to be relatively longer in small individuals than in larger individuals of a species. Gros and Cochard (1978) examined the relationship of abdomen length to carapace length (including the rostrum) in Nyctiphanes couchi; the carapace was shortest in mature females, longest in males and of intermediate length in immature females, relative to the length of the abdomen. Nemoto (1977) describes various biometrical relationships in Thysanopoda cornuta, T . egregia, T . spinicaudata and T . tricuspidata that can be used in the numerical taxonomy of these species. Other biometrical relationships are referred to in Mauchline and Fisher (1969). The growth rates of internal organs of the body relative to the body as a whole has only been examined in the case of the ovary of Meganyctiphanes norvegica (Mauchline and Fisher, 1969). Increase in the size of the ovary reflects, initially, an increase in the number of potential eggs and, secondly, the development of these eggs to a mature size. This process requires a period of approximately three months, as Hollingshead and Corey (1974) have confirmed for this species in Passamaquoddy Bay. Hollingshead and Corey also briefly describe the development of the testes and spermatozoa in this species and confirm that 8-10 weeks elapses between initial activity in the testes and production of fully formed spermatophores. Studies of the brood sizes of different species are described in Mauchline and Fisher (1969). Eggs are carried externally by the females in the genera Nyctiphanes, Pseudeuphausia,, Nematoscelis and Xtylocheiron while species in the other genera lay their eggs freely

10.

GROWTH, MATURITY AND MORTALITY

495

into the sea. Mauchline and Fisher include Nematobrachion and Tessarabrachion among the genera that protect their eggs by carrying them but this has not been observed and species in these genera probably lay their eggs directly into the sea. Direct counts of the number of eggs per brood can be made in females that carry eggs. Species that lay their eggs directly into the water can occasionally be induced to lay them in the laboratory but may not lay a normal sized brood under these conditions. Several authors have examined the numbers of eggs at a late stage of development in the ovary and estimated brood size by counting them (Table XVI). These eggs are usually at developmental Stage IV (Mauchline and Fisher, 1969). Mauchline found that the ripe ovary represented %lo% of the body volume of the female, while Roger (1973g, 1974c, 1978a) found that its weight ranged from 5.4 to 9.5y0, with a mean value of 8.2y0,of the female’s body weight in a study of nine species. The spent ovary of Meganyctiphanes norvegica has a volume of about 50% of that of the ripe ovary. Consequently, knowing the diameter and so the approximate volume of the newly laid egg, an estimate of brood size can be made. This estimate, of course, assumes that the egg does not swell significantly upon extrusion into the sea. Nemoto et al. (1972)found that the weight of the egg mass of Nematoscelis dificilis was, on average, 17.5% of that of the female’s body weight. This is surely indicative of swelling by intake of water. Consequently, Mauchline’s estimates, in Table XVI, under “other estimates”, are likely to be lower than the actual numbers of eggs laid. The available data on brood sizes of euphausiids have been summarized in Table XVI. Roger (1976) points out that closely related species have comparable brood sizes and so with the possible exception of the baChypelagic species in the genus Thysanopoda, the brood sizes of most species not listed in Table XVI can probably be deduced. The bathypelagic species tend to have small broods relative to their body size consisting of larger eggs (Mauchline, 1972); this has been confirmed by Nemoto et al. (1977) who estimate newly spawned eggs of T . cornuta to be 2.5-3.0 m m and those of T . egregia to be 0.82.0 mm in diameter and the brood size to be as indicated in Table XVI. There is still considerable conjecture about the brood size of the large Antarctic Euphausia superba. Everson (1977), quoting the work of Makarov, Naumov and Nemoto, points out that their estimates range from 2110-14 086 eggs per brood as compared with Mauchline’s estimate of 310-800. These high counts are derived from counting eggs within the developing ovaries and making assumptions about which of these eggs will be laid and which will not be laid. A considerable

TABLEXVI. BROOD SIZESOF EUPHAUSIIDS N u d e r of ripe eggs in Stage I V ovary Species

Body length (mm)

Bentheuphauaia amblyops Thysanopoda monacantha T . cristate T.tricuswa T . aequalis T . pectinata T . orientalis T . cornuta T . egregia Meganyctiphanes norvegica Nyctiphanm cowhi N . australis N . capensis N . simplex Pseudeuphawia sinica EuphauSia eximia E . krohni E. b r e d E . dhmedaw

25-35 25-30 35-45 15-22 17-19 29-40 25-38 70-90 45-60 19-36 10-15 13 13 15 9.5-12-5 22 12-18 7-10

E . super&

41-55 1s

E . luc6m-w

12-17

Range

Meen

1-7 32-72 20-76 20-82 16-31 30-82 38-74 86-198 191-265 350-556 50-60

3.5 54 40 40 24

Direct counts of eggs

54

54

50

20-96 ca 50 30-100 42-110 113

20-87 25-68

Other estimates

400-530 50 60 40 50 23-58 175

50 50 80

References

9 9 9 9 5, 9 9 9 11 11 3, 5 3, 5, 12 3 3, 6 3 3 3 5 5

50-200

7, 9 310-800 so

3 3

E. pacificen

E . nana E . gibba E . fallax E. hemigibba E . triacantha Thyaanoe8sa inermis T . longicaudatu T . gregaria T . rmchi Nematoacelis d i f i i l i s N . megalopa N . tenella N . athntiea Nematobrachion jlempa N . boopia Stylocheiron carinaturn S. afine S. suhrni S. microphthalm S.elongaturn. S. longicorne S. abbreviatum 8. maximum

IS16

5G100

18-24 8

50-200 10-12

12-15 32 26 13 8-1 1 26 22-25 15-25 15-21 12-14 22 19-2 1 11 7 7-9 7 13-16 7-10 16-19 20-30

35-43 100-230

60 80 60 310 300-350 325 160-172

150

215-400 168-305 50-80 163-205 12-39 4-2 1

300 200

300-400

49-66 74-101

180-405 220-250

65 150 19 9

50 80

62-143 170 3-14 4-12 5-15 10-13 2-5 4-16 13-22 50

8-10

3

3, 8 3, 8 3 9 9 9 3 3 3 5 3 2, 4 5, 10 9 5, 9 3, 9 9 1, 3, 6, 9 6 3, 5, 6 6 6 5 5, 9 5, 10

References: 1, Sebastian, 1966; 2, Komaki, 1967; 3, Mauchline and Fisher, 1969; 4, Nemoto etuZ., 1972; 5, Casanova, 1974; 6, Talbot, 1974; 7, Brinton, 1975; 8, Brinton, 1976; 9, Roger, 1976; 10, Mauchline, unpublished; 11, Nemoto et ul., 1977; 12, Gros and Cochard, 1978.

498

THE BIOLOGY OF EUPIIAUSIIDS

number of the developing eggs within the ovary never mature but are resorbed after the brood has been produced. The majority of estimates of the sizes of the large broods quoted above are within a range which is equivalent to 30-40% of the body volume of the female parent. Such broods would be exceptional in size not only among euphausiids (Table XVI) but among crustaceans in general where brood size is normally within the limits of 5-10y0 of body volume. Consequently, as Everson points out, further investigation of the brood size of this species is urgently required. Brood size of a species increases with increasing body size (Mauchline and Fisher, 1969). This is illustxated in Euphausia pacijica,Nematoscelis megalops and N . dificilis (Fig. 23). Brinton (1976) counted Stage I V eggs in the ovaries of Euphausia pacijica of known body length and found disproportionately small numbers of such eggs in

500 -

-

200 v)

% B L

W

100-

-

-

a

5

Nematoscehs difficihs

2

50 -

Euphausia pacifica

200

20-

lo4

103

Body length3

50

100 Body weight (mg)

200

FIG.23. Brood size related to body size. Left: numbers of eggs per egg mass of Nematoscelis megalopa in the north-east Atlantic related t o body lengthS (Mauchline, unpublished) compared with the numbers of Stage IV eggs in the ovaries of EuphazLeio paci$ca off California (Brinton, 1976). Right: numbers of eggs per egg mess of Newtoscelis di&ilL in the north-west Pacific related to body weight; dots sfter Komaki (1967), triangles after Nemoto et al. (1972).

10.

GROWTH, MATURITY AND MORTALITY

499

the ovaries of the largest size class, 20-21 mm total length. Samples of Nemotoscelis megalops collected by R.R.S. “Challenger” near 55”N, 12”W in May-Julg, 1973 and 1975 were examined and the number of eggs per brood counted (Fig. 23). The slopes of the two lines are closely similar. Komaki (1967) relates the numbers of eggs per brood to the females’ body weight in N . dificilis and Nemoto et al. (1972) produce a similar relationship with fewer data. Gros and Cochard (1978) examine the number of eggs per brood in Nyctiphanes couchi and find that it increases with body size although there is a wide range of variation. There is a distinct tendency for more eggs to be present in the left hand egg mass than in the right hand one. Nemoto et al. (1972) describe the relationship of brood weight t o body dry weight as well as wet weight in N . diitficilis. The brood weight ranged from 13.3-24.4%, mean 17.5%, of the female’s wet weight. The corresponding values on a dry weight basis were 17.826.4%, mean 22.7%, of the female’s dry weight; this suggests that the eggs contain a smaller portion of volatile components than the female. They further found that the weight of individual eggs in the broods was greater in broods with larger numbers than smaller numbers of eggs. This means that larger females do not only produce larger numbers of eggs per brood but that the eggs themselves are also larger. This was not confirmed in broods of N . megalops from the north-east Atlantic (Mauchline, unpublished). There is still little further information available about the number of successive broods that an euphausiid can produce. Roger (1976) points out that %hetropical species that he examined do not release eggs continuously from an individual ovary, as suggested by Mauchline and Fisher. The females appear to release all ripe eggs at once or within a short period of a few days. There are then only very small eggs left in the ovary and a period of several weeks would be required for their development to a ripe stat,e. Thus, fecundity of a species cannot be discussed with confidence until the number of broods per female is known. Species in middle and high latitudes probably produce a single brood. This appears true in Meganyctiphunes norvegica. Thysanobsa raschi produces one brood in the spring but may, in some sea areas, produce a second brood in the autumn. Euphausia superba and Thysanoessa gregaria also probably produce one brood per breeding season (Makarov, 1975b). Three of these four species, however, have a longevity of 2-3 years and may breed in two successive years. It is probable that fecundity equals approximately 2 x brood size, but in small tropical or subtropical species it may be equal to brood size.

500

THE BIOLOGY OF EUPHAUSIIDS

Rates of growth of larval and adult euphausiids are described in Mauchline and Fisher. The rates were primarily estimated through statistical analyses of samples of wild populations, although Lasker's work on the growth and moulting of Euphausia paci$ca in the laboratory is included. More experimental observations of growth and moulting are now available and an empirical approach to analysing growth rates of crustaceans has been developed. Various significant linear correlations between the duration of the intermoult period and the growth factors at moulting on the one hand and body length, weight and successive moult number on the other have been defined in decapod crustaceans other than euphausiids (Mauchl+e, 1976, 1977a). The growth factor at moulting is defined as the increase in body length or body weight at moulting expressed as a percentage of the premoult body length or weight. The duration of the intermoult period was found to increase logarithmically, the growth factors to decrease logarithmically, when plotted against body length or successive moult number, illustrated for Euphausia supberba in Fig. 24 (p. 504). These relationships were examined in euphausiids by Mauchline (1977~)using the data of Fowler et al. (1971a' c), Gopalakrishnan (1973) and Le Roux (1973a). Log/linear regression equations relating the duration of the intermoult period to successive moult numbers or body length were calculated for the data on Nyctiphanes couchi, Nematoscebis dificilis and Meganyctiphanes norvegica ; the correlation coefficients were significant a t the 1 4 % level in eight of the ten equations. Sameoto (1976b) has since produced further information on the intermoult periods of M . norvegica and Thysanoessa inermis. He found that their durations, in each species, were linearly related to temperature over the range 0°-16"C. His data along with previous estimates of the durations of intermoult periods of other species are summarized in Table XVII. The only experimental data relating growth fac$ors to body length are those of Fowler et al. (1971a) on growth increments in length of Meganyctiphanes norvegica. There was considerable variation in the data and no significant correlation was present although the growth factors tended to decrease with increasing body length. The logarithmic decay of growth factors against increasing body length has been well enough demonstrated in other crustaceans (Mauchlme, 1977a) to allow its assumption in euphausiids. Hiatt found two inflexions when relating post-moult t o pre-moult carapace width in the crab Pachygrapsus crassipes and considered them associated with the change from larval to post-larval growth rates and with the attainment of sexual maturity (Mauchline, 1976).

10.

QROWTH, MATURITY AND MORTALITY

501

TBLE XVII. DURATIONS OF THE INTERMOULT PERIODS OF EUPHAUSIIDS AT DIFFERENT ENVIRONMENTAL TEMPERATURES

Species Meqanyctiphanes norveqica Nyctiphanes couchi N . simplex Euphausia eximia E . krohni E . pacifcca

E . superba Thysanobsa inspinata T . spinijera T . inemis T . raschi Tessarabrachion oculatum Nenzatoscelis megalops

T"C 4 10 13 13 8.3-12.2 11-17.5 14.8-16.4 13 6.5-16 8.3-12.2 9.5-18.9 12-15 14.8-16.4 1.4-3.5 11-15 8.3-12.2 11-15 0 10 11-15 11-15 13

Intermoult period (days)

Reference

13-19 4-11 7-9 4-10 5-7 4-6 3-4 4-5 3-8 5-6 4-7 4-6 3-4 12-17 6 5-6 5 15-16 6 6 5 5-7

Sameoto (197613) Sameoto (197613) Fowler et al. (1971a) Fowler et al. (1971a) Jerde and Lasker (1966) Jerde and Lasker (1966) Lasker and Theilacker (1965) Fowler et d.(1971a) Lasker (1966) Jerde and Lasker (1966) Laskr (1964) Paranjape (1967) Laaker and Theilacker (1965) Mackintosh (1967) Paranjape (1967) Jerde and Laaker (1966) Paranjape (1967) Sameoto (197613) Sameoto (197613) Paranjape (1967) Paranjape (1967) Fowler et al. (1971a)

The first inflexion appears to be real, the growth factors of successive larval stages tending to decay at a, different and usually faster rate than those of the post-larval stages. The second inflexion may be an artifact originating from the fitting of a line rather than a curve to his data. There is, however, some evidence that growth factors of crustaceans may change once sexual maturity is attained. Some decapods do not have an increase in body length at the moult immediately before the eggs are laid but a normal increment is achieved at the following moult. This situation would produce an inflexion. The growth of the calyptopes of 27 species of euphausiids has been examined. The following authors provide measurements of the calyp topes and first furcilia so that the growth factors a t each of the successive moults can be calculated: F'raser, 1936; Einarsson, 1945 ; Heegaard, 1948; Boden, 1950, 1951, 1955; Sheard, 1953; Lewis, 1955; Bary, 1956; Baker, 1959; Mauchline, 1959, 1965, 1971b; Wang, 1965; Mathew, 1971, 1972; Gopalakrishnan, 1973; Knight, 1973, 1975, 1976;

502

THE BIOLOGY O F EUPHAUSIIDS

and Casanova, 1974. The growth factors were found 130 fall into two natural groups (Table XVIII). Species in the genera Nematoscelis and Stylocheiron have lower growth factors than those in other genera. Allometric growth of the abdomen takes place in the calyptopes but such growth is insignificant in the furciliae, the body proportions remaining relatively constant (Sheard, 1953). The growth factor in body length of 26+8% of the CIII-FI moult of the majority of euphausiids (Table XVIII) is equivalent to a doubling of the body weight at this moult. TABLE XVIII. MEAN PERCENTAGE GROWTH FACTORS AT MOULTING OF CALYPTOPES.SPECIESIN GROUP B BELONG TO THE GENERA Nematoscelis A N D Stylocbiron, THOSEIN GROUPA TO TRE OTHER GENERA Number of species

Group A Group B

23 8

CI-CII 5 4 k 17 37 29

+

CII-CIII

CIII-FI

3 6 2 10 25+9

26+8 11$7

C , calyptopis. F, furcilia.

The growth factors of the successive furciliae of 28 species were examined, the data being obtained from the references listed above for measurements of the calyptopes with the addition of John (1936), Brinton (1962) and Jones (1969). Log/linear regression equations relating growth factors to successive moult number were calculated and the correlation coefficients found t o be significant at the 1-5% level in 15 of the 28 species (Table XIX). The least degree of correlation was found in the 6 species examined in the genera Nematoscelie and St ylocheiron . The development of the larvae of euphausiids, especially species living in coastal areas, is variable. I n a species, larvae of the same body form can vary in body length at different times in the same sea area or a t the same time in different sea areas (Mauchline and Fisher, 1969). This is also evident in the data on Thysanoesssa Zongicaudata where Jones (1969) found that the larvae in a warm sea area were smaller than the larvae in a cold area off Newfoundland. This caum difficulties in estimating the mean body length of any one larval stage because samples of a species for such investigations are often taken over wide geographical areas, and sometimes over more than one breeding season. Baker (1959) and John (1936), for example, both

TABLEXIX. REGRESSION ANALYSES OF GROWTH FACTORS ON SUCCESSIVE MOULT NUMBERSOF FURCILIAE

Species

Number of moulu

Thysanopoda acutifrons Meganyctiphtcnes norvegica Nyctiphanes couchi N . australis N . simplex Pseudeuphuaia sinica Euphausia krohni E . diomedeae E. superba E. vallentini E. lucens E. frigida E . pacifica E. w n a E. sibogae E. fallax E. gibboides E. sanzoi E . triacantha ThysanoEssa inemnis T . longicaudata T . raechi Nematoscelis megalops N . dificilis field laboratory

Stylocheiron wuhmi S. carinatum 8.longicorne 8.abbreviatum

Regression constants log y = bx + a a b

Value an,d significance of 'p

Source of original data

7

1.54

-0.12

-0.937**

Einarsson (1945)

7

1.45

- 0.03

- 0.540

Mauchline (1959)

6

1.58

- 0.12

- 0.866*

Casanova (1974)

6 4 8

1.55

1.51 1.52

- 0.10 - 0.09 - 0.07

- 0.878* - 0.720

Sheard (1953) Boden (1951) Wang (1965)

5 6 6 6 5 6 6 4 6 6 6 6 6 6 9

1.45 1.32 1.42 1.76 1.72 1.82 1.51 1.60 1.50 1.46 1.36 1.39 1.89 1.64 1.29

-0.11 - 0.07 - 0.05 - 0.17 -0.16 - 0.17 - 0.10 - 0.14 -0.16 - 0.12 - 0.06 - 0.08 - 0.18 -0.10 - 0.05

- 0.794

7 5 11 5

1.39 1.48 1.32 1.36

- 0.08 - 0.09 - 0.03 - 0.04

- 0.782* - 0.885*

- 0.581

4

1.47

- 0.22

- 0.835

6

1.30

- 0.07

- 0.614

5 6 4 7

0.98 1-14 1-33 0-93

- 0,004 - 0.05 -0-11 - 0.002

- 0'082 - 0.795 - 0.451 - 0.008

-0.916**

- 0.673

- 0.705 - 0'702 - 0.948* - 0'979*** - 0*964** - 0.718

- 0.912*

- 0.954** - 0.858* - 0.077

- 0.905*

- 0.938** - 0.995***

- 0.603

Casanova (1974) Mathew (1971) Fraser (1936) John (1936) Bary (1956) John (1936) Boden (1950) Brinton (1962) Mathew (1971) Knight (1978) Knight (1975) Knight (1976) Baker (1959) John (1936) Einarsson (1945) Einarsson (1945) Jones (1969) Mauohline (1965) Casanova (1974)

Gopalakrishnan (1973) Gopalakrishnan (1973) Casanova (1974) Mathew (1972) Casanova (1974) Casanova (1974)

504

THE BIOLOGY OF EUPHAUSIIDS

used samples from a wide area of the Antarctic, John’s samples being collected over several seasons in the 1930s, Baker’s samples being also collecbed in the 1930s and combined with others taken more recently. Consequently, some of these practices may account for the fact that significant correlations were only found in 15 of the 22 species examined in the genera Thysanopoda, Meganyctiphanes, Nyctiphanes, Pseudeuphausia, Euphausia and Thysanoessa. A growth factor line and intermoult period line, as illustrated in Fig. 24, can be determined for a species by using the following minimum of data : (a) The time in days required for a known number of moults or for a few individual moults where the corresponding body 40

7l 5

Larvae

-

1

I I I 1

I

Adolescents

Adults

I I

I

I

1

I

I

I

FIG.24. The intermoult period and growth factor lines for Euphawria superba. The growth factors of the larvae decay at a faster rate than those of the adolescents and adults. The regression equations of the three lines are given in the text.

lengths are known. These data can be obtained in the laboratory or by sampling natural populations especially larvae and early post-larvae. An accurate estimate of the increments in body length eb moulting for a few successive moults or for a few moults where the appropriate pre-moult lengths are recorded; or an accurate estimate of the increase in body length over a known number of moults. An approximate growth curve for the animal throughout its life, derived from seasonal sampling of natural populations. attempt has been made t o reproduce such lines to describe the growth of Euphausia superba. Mauchline (1977~)produced a preliminary study but some further data are now available. There are three

10.

GROWTH, MATURITY AND MORTALITY

505

bits of information. available on the intermoult periods of E . superba that can be used t o determine the intermoult period line. Firstly, Mackintosh (1972) suggests that the egg and subsequent larval stages may each have durations that are about twice as long as those of the comparable stages of Meganyctiphnes norvegica. Furciliae of M . norvegica moulted in the laboratory every 4-7 days and so comparable stages of Euphausia superba may moult every 8-14 days in the lower temperature regimes of the Antarctic. Secondly, Fraser (1936) states that the development of the six furciliae of E . superba is not completed by the end of April but that the larvae are then entering furcilia V. This is confirmed by Makarov (1974b) who found tihat the larval development was not completed until the following October. Mackintosh (1972) concludes that the eggs are spawned at the end of January and consequently development from egg to the furcilia I V at the end of April, ten moults in all, would take place in a period of about 90 days. Thirdly, Mackintosh (1967) concludes from experimental observations that the intermoult periods of individuals of body length 20-33 mm range from 12 t o 17 days. Clarke (1976) has confirmed this range of duration. These data satisfy the regression equation

log IP = 0*9589+ 0.009OL

(1)

where I P is the intermoult period in days and L is body length in mm. The successive intermoult periods are calculated in Table XX : furciliae moult every 10-12 days; 94.4 days are required for the egg to develop to the furcilia I V and individuals of body length 20-30mm have intermoult periods of 14-17 days (Fig. 24). A growth factor line for the larvae is easily derived from the data of Fraser (1936). He examined the larvae from samples taken over a wide geographical area of the Antarctic Ocean and in different years, and calculated the mean body lengths of the different stages. The third calyptopis increases its body length by 26% when it moults to the furcilia I. Fraser’s data satisfy the regression equation log Gf = 1.6099- 0.0503 L

(2)

where Cf is the larval growth factor and L is body length in mm. This equation, as would be expected, is slightly different from that for the regression of growth factors on successive moult number given for this species in Table XIX. There is no information available on the growth factors a t successive moults of adolescent and adult E . superba except those of McWhinnie et al. (1976) who state, as a result of rearing experiments, thati “the

TABLEXX. HYPOTHETICAL LIFEHISTORYOF Euphausia superba Dates of moulta in annual growing period of Moult No.

Stage

Body length (mm)

Growth Intermoult Cumulative factor period growing (yo) (days) period

180 days

8.14 16.61 25.39 34.47 43.83 53.46 63.34 73.44 83.80 94.40 105.29

23.i 5.ii 13.ii 22.ii 3.iii 12.iii 22.iii l.iv l0.iv 20.iv 1.v 1l.v

150 days

120 days

4.00 5.04 6.18 7.41 8.69 9.98

26.00 22.72 19.90 17.27 14.89 12.81

8.14 8.47 8.78 9.08 9.36 9-63 9.88 10.10 10.36 10.60 10.89 11.19

11.26 12.51 13.94 15.47 17.11 18.85 20.68 22.60 24.60 26.67 28.80

11.06 11.40 10.99 10.58 10.15 9.72 9.28 8.85 8.41 7.99 7.57

11.49 11.79 12.14 12.53 12.97 13.45 13.96 14.53 15.15 15.81 16.52

116.48 127.97 139.76 151.90 164.43 177.40 190.85 204.81 219.34 234.49 250.30

First winter 1.ix 13.xi 25.xi 7.xii 2O.xii 3.i 17.i 31.i 16.ii 3.iii

23 24

30.98 33.20

7.17 6.78

17.29 18.10

266.82 284.1 1

25

35.45

6.41

18.97

302.21

2O.iii ___ 1.xi 6.iv - 18.ix 24.iv l.xi 6.ii __

26 27 28 29

37.72 40.00 42.29 44.57

6.05 5.72 5.40 5.09

19.88 20.84 21.85 22.91

321.18 341.06 361.90 383.75

1.xi 21.xi 12.xii 3.i

30 31 32

46.84 49.09 51.32

4.81 4.55 4.30

24.01 25.16 26.35

406.66 430.67 455.83

13.ii l.xi 25.i 18.ii 9.iii 25.xi 4.iv 2O.xii 16.iii __

33 34

53.53 55.71

4.07 3.85

27.58 28.86

482.18 509.76

35

57.85

3.65

30.17

538.62

1l.iv l.xi 16.1 9-v 29.xi 1 2 3 1.xi 27.xii 13.iii

36 37

59.96 62.03

3.46

31.52

568.79 600.31

Egg 1

2 3 4 5 6 7

8 9 10 11 12 13 14 15 16 17 18 19 20 21 22

Egg NI NII Mn CI CII CIII FI FII FIII FIV FV FVI Adol.

-

~

~

2O.xi 25.xii l0.xii 14.i 3l.xii 4.ii 22.i 26.ii

-

~

~

~ ~

1.xii 27.i 2.i 27.ii

1.xi 2.xii

The dates at which different moults would take place in life cycles with summer 8888on8 of active growth of 180, 150 and 120 days are shown assuming that eggs are laid on 28 January. The double lines across the columns indicate periods of little growth in the winter. Further details are given in the text.

10.

GROWTH, MATURITY AND MORTALITY

507

growth increment per moult of juvenile to young adults ranged from 0 to 10 percent of the previous intermoult size”. The growth factors of decapod crustaceans decay throughout the life of the animal such that they are usually in the range 2-6% at sexual maturity (Mauchline, 1977a). The growth factors described by the regression equation for the larvae (equation 2) are decaying a t too fast a rate t o describe growth of the adults. Using this equation, a growth factor of 4% occurs in individuals of body length 20 mm, a factor of 1% in animals of length 32 mm and a factor of 0.1% in animals of 52 mm, as can be extrapolated from Fig. 24. Thus, the animal would take many years to attain its adult size if its growth factors are described by this equation. An inflection in the growth factor lineis often present a t the end of the larval development of crustaceans (Mauchline, 1976, 1977~). Thus, the only way of fitting a provisional growth factor line to the adults is to assume that, like other decapod crustaceans, the growth factors will decay from the values present a t the end of the larval development to values of between 2% and 6% in individuals tha.t have atkained sexual maturity. Sexual maturity is achieved at a body length of over 40mm and so the growth factors would be expected to have decayed below 6% by that time. The regression equation log Gf = 1.1930-0*0109 L

(3)

where Gf is the growth factor and L is body length satisfies this condition, an animal of 40 mm length having a growth factor of 5.7% (Fig. 24). The slope of this line is also governed to some extenb by the slope of the intermoult period line in the region of animals of length 20-30mm in order to satisfy the observations of Mackintosh and Clarke on the intermoult periods corresponding to animals of these lengths. A hypothetical life history can now be constructed (Table XX). The third calyptopis (CIII) has a body of length 4 m m . Its growth factor and intermoult period can be calculated from regression equations (1) and (2). Application of the calculated growth factor t o the body length of the C I I I produces the body length of the fist furcilia (FI).The process is repeated with the body length of the FI to calculate its intermoult period, growth factor and so the body length of the FII. This process continues to the first postlarval stage when equation (3) is substituted for equation (2). Then, using equations ( I ) and (3), the calculations are continued for the post-larval stages. Calculaeion of the growth factors of larval stages earlier than the CIII is not valid because allometric growth is present in the developmental

508

THE BIOLOGY O F EUPHAUSIIDS

sequence nauplii-metanauplius-CI-CII-CIII. Calculation of the intermoult periods of these larvae has t o be made by transforming the regression of intermoult period on body length to a regression of intermoult period on successive moult number. The cumulative growing period (Table XX) shows $hat the equations describe an animal in which the egg develops to the F I V in a period of 94.4 days. The equations also state that the animal requires a total of 600 days of continuous active growth to reach a body length of 62mm, the approximate maximum body length recorded. Active growth in length, however, will not be a continuous, non-fluctuating process in E. superba. Animals in higher latitudes tend to have a winter period during which increase in body size is minimal or completely absent. This has been demonsbrated in many decapods and also in other species of euphausiids (Mauchline and Fisher, 1969). There is no evidence to suggest that E. superba will be different and confirmation of this was obfained by Clarke (1976), who found an increase in the intermoult period associated with a decrease in the water temperature in his experimental containers. No moulting took place in the krill that survived into winter. As stated earlier, Fraser (1936) believed that the larval development ceased a t the end of April, further moulting being inhibited. Mackintosh (1972) estimated the rate of growth in several regions of the Antarctic Ocean, his curve of growth rate in the Northern Weddell Drift being supported by the most data. The growth curves drawn in his Figs 44 and 45 indicate long seasons of active growth; they extend from the beginning of September through to April or May, some 240-270 days in length (Fig. 25). The northern euphausiids, subject t o marked seasonal variations in environmental temperature, have growing seasons of 150-180 days and it is unlikely that the growing seasons of E . superba, living in higher and colder latitudes, will exceed trhis. As stated earlier, Clarke (1976) found evidence of inhibition of moulting by winter temperatures in this species, and Makarov (1971a) suggests that the seasonal period of active growth in length begins significantly earlier in warmer years. The determination of the duration of the seasonal period of active g r o d h is central t o this method of analysing growth of a crustacean. Hart (1942) shows that increased populations of phytoplankton are present from about the beginning of November, being maximal in the period December to February, and extending into May. This is a period of about 180 days. Moulting, and therefore active growth, may, however, be more closely controlled by environmental temperature than the presence of phytoplankton.

10.

60

509

GROWTH, MATURITY AND MORTALITY

-

l8Oday growth period

-

150 day growth period

50

-

Fits curve of Mackintosh-

-

40

E

E

120 day growth period

-

1

I$

300

m

/Y

20

-

F5

I I

I I I I F

A

J

A

I

I

I

I I

I Year ‘2

I

I

Year 3

I

I

1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 1 0 D F A J A 0 D I F A

Year4

I I I

I

I I I l l l I J A 0 D

1 1

I I I I I I

F

A

J

A

Months

FIG.26. Theoretical growth curves derived from the regression equations in the menner shown in Table I X . The open circles are values from the growth curve deduced from seesonal population histograms by Mackintosh (1972) ; a theoretical curve, solid circles, is fitted to these values. The other curves are from Table XX and represent patterns in populations with summer periods of active growth lasting for 180, 150 and 120 days respectively. F5 is the fifth furcilia.

The data in Table XX have been organized to describe annual growth in terms of 180, 150 and 120 day summer periods of active increase in body length. The corresponding curves are illustrated in Fig. 25. For the sake of easy comparison, seasonal active growth is considered to commence on 1 November and continue for the required number of days.

510

THE BIOLOQY OF EUPHAUSIIDS

The curves are based on the patterns of growth of northern species of euphausiids and of decapods in general and incorporate the tentative available data on growth and moulting of E . swerba. The curves with 180 and 150 day annual growth periods suggest thab sexual maturity, attained at lengths of 40-50mm, is reached a t an age of two years. The curves also suggest that animals of body length greater than about 55 rnm are approaching an age of three years. This is the situation described by Mackintosh in the warmer Northern Weddell Drift. The curve with a 120 day annual growth period still suggests that part of the population achieves sexual maturity at two years of age, although a portion of it may not mature until the following year. This curve suggests that the largest recorded animals would then be approaching four years of age. A reduction in the annual period of active growth to 90 days would require most animals to be three years old before reaching sexual maturity and five years old a t a length of 60mm. This latter situation describes the conclusions of Ivanov (1970) in the colder East Wind Zones. Samples of krill have been collected and analysed since the 1920s but growth rate and longevity are still not known (Everson, 1977; Kock and Stein, 1978). There is no evidence to suggest that similar analyses will be more successful in the future. The present approach is different and may produce information that can be used to re-interpret the data already available. It requires repetition of the experiments and observations of Mackintosh (1967), Clarke (1976) and McWhinnie et al. (1976) who maintained E . superba on board ship and observed it moulting. Data, obtained on the durations of the intermoult periods and the growth factors a t moulting of krill of different sizes from different regions can be used $0 test and provide more accurate equations than those presented here. Observations of the effects of temperature and food upon moulting can help in defining the duration of the seasonal period of active growth. The revised equations can in turn be used to reinterpret the infcmnation derived from length/frequency histograms of populations. The available information on the age and body length at sexual maturity, number of successive breeding seasons, and longeviky is summarized in Table XXI. Many of the species investigated mature a t one year of age, breed once and the majority of the population die shortly afterwards. Species such as Meganyctiphanes norvegica mature a t one year of age, breed and a major portion of the population survives to breed for a second time a t an age of two years. A few species have been shown to mature sexually at an age of two years and then breed for the first time. Some populations of Ewphawia superba may not

TAELEXXI. MATURATIONAND BREEDING OF EUPHAUSIIDS

Species

No.of M&mum Age at Size at years life nzaturity mturity breeding expectancy

1 Thysanopoda tricuspidata T. acutifrons 2 T. monacantha 1 T . aequalis 1 T. cornuta T . egregia T . spinicaudatu Meganyctiphanes norvegica 1

Nyctiphanes couchi Euphausia krohni E . diornedeae E . superba E . paci$fica E . triacantha Thysanoasa spinifera T. longipes T. inermis

T . longicaudata T . gregaria T . raschi Nemztoscelis megalops N . tenella Stylochiron longicorne

1 1 1 2(?) 1 2 2 1-2 1 2 1 or less 1 1 2 1 1 1-

> 25 > 35 > 25 > 16

Breeding smon

1 1 1 1

1 2+ 1+ 1

All year May All year All year

2

3+

Spring/Summer

2 1 1 2 1 2 1 1-2 1-2 1-2 1 (9) 1 2 1-2 2(9) 1 1

2 1 1 3 1 3 2 1-3 1-2 2-3 2 (9) 1 2+ 2-3 2)9~ 1 1

February-August May-August All year November-April All Year October-November June-September MayJuly MarchJune AprilJune April-October

ca 70 40-60 80-90 > 22

> 12 > 14 > 15 > 45 > 11 > 30 > 20 > 14 > 14 > 22 > 10 >8 > 14 > 22 > 17 > 16

>6

+

+

+

+

MarchJune May-August FebruaryJuly All year

Area

References

Eq. Pacific N. Atlantic Eq. Pacific Eq. Pacific

10 1, 13 10 10 18 18 18 N. Atlantic 1, 3, 7, 9, 11, 12, 14 G. of Gascogne 20 N. Atlantic 3, 11 Eq. Pacific 5, 10 Antarctic 1, 4, 8, 21 N. Pacific 1, 6, 15, 19 Antarctic 1 N. Aleutians 1 N. Pacific 1, 16, 19 N. Atlantic 1, 3, 7, 13, 14, 23 High latitudes 1, 2, 19 N. Atlantic 1, 3, 14, 22 N. Atlantic 3 N. Atlantic 1, 3, 7, 14, 17 High Latitudes 1, 19 Norway 11, 13 Eq. Pacific 10 Mediterranean 11

References: 1. Mauchline and Fisher, 1969; 2, Geiger et al., 1968; 3, Jones, 1969; 4, Ivenov, 1970; 5, Weigmann, 1970; 6, Smiles and Pearcy, 1971; 7, Wiborg, 1971; 8, Mackintosh, 1972; 9, Matthews, 1973; 10, Roger, 1973g, 1974c, 19788; 11, Casanova, 1974; 12, Hollingshead and Corey, 1974; 13, Jorgensen and Matthem, 1975; 14, Berkes, 1976; 15, Brinton, 1976; 16, Zhuravlev, 19768; 17, Berkes, 19778; 18, Nemoto et al., 1977; 19. Shvetsova, 1977; 20, Gros and Cocherd, 1978; 21, Kock and Stein, 1978; 22, Lindley, 19788; 23, Kulke and Corey, 1978.

512

THE BIOLOGY OF EUPHAUSIIDS

mature until at least three years old and this is probably also true of some of the mesopelagic and bathypelagic species. A few species are now known to have markedly shorter life histories than any of those described above. Recent investigations have shown that populations of euphausiids, not only in equatorial regions but also in middle latitudes, as instanced by Euphausia pacijca off California and Oregon (Smiles and Pearcy, 1971; Brinton, 1976), have a proportion of breeding females at all times of the year. There axe, however, periods of more intense breeding; the timing of these periods of intense breeding may vary from year to year and can, as off California, be related to events such as those associated with upwelling (Brinton, 197 6). Consequently, several overlapping generations may be produced in these populations during the course of the year. The life expectancy or longevity of individuals can differ from generation to generation. Euphausia pacific& has a maximum life expectancy of about one year in these regions. Individuals produced early in the year reach maximal size in a shorter period, about 6-8 months, than those produced later in the year, about 12-14 months. There is a great similarity between the structural elements of these populations of E . paciftca and those of populations of temperate species of Mysidacea described elsewhere in this volume. The earlier work of Nemoto in the north-western Pacific shows that seasonaliby of breeding was quite marked and there was no suggestion of continuous breeding throughout the year (Mauchline and Fisher, 1969). A different situation has been described in populations of Thysanoessa raschi and T . longicaudata. The former species sometimes has two breeding seasons in the Firth of Clyde, Scotland, one in the spring, the other in hhe early autumn (Mauchlinednd Fisher, 1969). This may be an example of the same adults maturing twice in the same year to produce two broods or it may result from the presence of two cohorts within the population as occurs in Nyctiphanes cowhi in the Gulf of Gascogne. Gros and Cochard (1978) found that there is continuous breeding within the population of this species throughout the period February to August. There are, however, two periods of intense breeding, the first a t the end of the winter, the second in the summer. These periods result from two cohorts within the population that breed a t separate times. A small proportion of each cohort survives a second winter and may breed for a second time in the respective season of the following year. Thysanoessa longicaudata, on the other hand, produces discrete generations in some regions of the eastern North Atlantic, the spring generations maturing sexually to

10.

GROWTH, MATURITY AND MORTALITY

513

breed in the autumn (Lindley, 1978a). The autumn generation forms the overwintering population that matures sexually the following spring. The only other indication of the production of multiple generations is in populations of Nyctiphunes australis in Hauraki Gulf, New Zealand; Jillett (1971) observed three breeding periods in these populations. This recent work has demonstrated a range of breeding patterns in euphausiids varying not only between species and latitudes but also within the same species in different regions or at different times in the same region. This brings us back to the problem of the life history of Euphausia superba. It reinforces the suggestion that the breeding patterns may vary in successive years, making the interpretation of length/frequency histograms difficult. There is still no hard information on the growth rates, breeding and longevity of the bathypelagic species of the genus Thysanopoda and of Bentheuphausia amblyops. The numbers of young per brood relative to body size appear to be fewer than in epipelagic species (Table XVI, p. 496). Nemoto et al. (1977) have re-examined the body length a t sexual maturity in the three Thysanopoda species (Table XXI). These lengths are approximately the same as reported previously and do not alter the tentative conclusions about the life histories of these species drawn by Mauchline (1972). The growth rates of high latitude epipelagic species were extrapolated to body lengths of these large bathypelagic species. These extrapolations suggesh that the bathypelagic species probably live 3-7 times as long as epipelagic species and do not mature sexually until an age of at least two years. The larger species, T . spinicaudata and T . cornuta, probably do not mature until their third or fourth year. The maximum life expectancy of T . spinicaudata may be in the range 12-20 years. Mauchline and Fisher (1969) briefly discuss the apparent influence of environmental temperature on the maximum size attained by different species. The influence of temperature may not be direct but, of course, act through consequent availability of food. Geiger et al. ( 1968) present interesting length/frequency histograms of populations of Thysanoessa inermis in the Arctic Basin and off north-east Greenland. The Greenland population is bimodal but the population in the colder Arctic Basin only has a single mode corresponding to the smaller size mode of the Greenland population. They recorded one male and 84 females in the Arctic Basin samples and conclude that this species was probably not breeding there. Changes in the sex ratios in populations of euphausiids have been found to occur at the times of spermatophore transference from the

514

THE BIOLOGY OF EUPHAUSlTDS

males to the females and also during the period of egg laying by the females (Mauchline and Fisher, 1969). Wiborg (1971) found such changes in the sex ratio at these times in populations of Meganyctiphanes norvegica, Thysanoessa inermis asd T . raschi in the Byfjord and Hardangerfjord, Norway. Sex ratios ip populations of Meganyctiphanes norvegica in Passamaquoddy Bay were variable but a similar general pattern of change was observed by Hollingshead and Corey (1974). Ratios of males to females varied from 0.1-1.7 in populations of Thysarw2irsa inermis in the Korsfjorden, Norway (Jorgensen and Matthews, 1975). Baker (1970) examined a surface swarm of Euphausia krohni off Fuertaventura, Canary Islands and found a male t o female ratio of 1.1 : 1.0 which compared with a ratio of 0.78 : 1.0 in normal net samples of this species. Sebastian (1966) found a comparable ratio of 5 : 1 in a shoal of Thysanopoda tricuspidata a t 8'10'N, 75"55'E in the Indian Ocean. Sex ratios in populations of Euphuusia pacijca off California have been examined in some detail by Brinton (1976). He has shown that bhe ratio of females to males increases with size because females have either a lower rate of mortality or they grow a% differentr rates from the males. Breeding is a more or less continuous process in the Californian populations and so no changes in sex ratio could be correlated with spermatophore transference or egg laying. Nemoto et al. (1977) state thab females normally outnumber males in populations of epipelagic species. The male to female ratio in the majority of species throughout most of the year is probably in the range between unity and 0.7 : 1.0. Females attain a larger body size than males in many species through either increased growth raDes, increased longevity or a combination of both. Consequently, the sex ratio is very much in favour of females among the large size classes of a species.

CHAPTER 11

ECOLOGY OF DISTRIBUTION The distributions of all the species of euphausiids throughout the oceans are described in detail in Mauchline and Fisher (1969) and supplementary information is given in an earlier part of this review. The ecological aspects of the distributions were discussed against a background of the general topographical and physical-chemical features of the oceans. Little further information can be usefully added to those general descriptions but additional studies of various regional faunas have now been made. Several of these regions have not been previously investigated in any detail. The most significant and largest is of course the Indian Ocean which was intensively sampled during the International Indian Ocean Expedition 1960-1965. The distributions of many species of euphausiids are related to such features as gyses and transition zones between water masses. McGowan (1974, 1977) and Reid (1977) examine the structures of ecosystems and the features of the boundaries between them. Much of t,his work is concerned with the Pacific Ocean where the distributions of the faunas are better known. McGowan (1974) illustrates the distributional patterns of the basic biotic provinces of the Pacific and Reid (1977), using species of euphausiids as examples, discusses the distributions in relation t o the physical-chemical features of the environment. The partitioning of the different species between the different oceanic regions is given for each genus in Mauchline and Fisher (1969). Gopalakrishnan (1974) has made a detailed study of the zoogeography of the seven species in the genus Nematoscelis. The euphausiid fauna of the different regions of the oceans are described in Mauchline and Fisher and the following additional information is presented here under the same regional headings, except for the “Arctic Ocean”, a heading not used in the previous work.

Arctic Ocean and peripheral seas Few species occur in the Arctic Ocean and only Thysanoessa raschi and T . inermis are common (Geiger et al., 1968). Two boreal species,

516

THE BIOLOGY OF EUPHAUSIIDS

T . longipes and T . longicaudata, were also found but only rarely. Peripheral regions of the Arctic Ocean are penetrated by North Atlantic and North Pacific water masses transporting these latter two species as well as Meganyctiphanes norvegica, Nematoscelis megalops and, very occasionally, Stylocheiron maximum. Tile euphausiid fauna of the Barents Sea is well known and Mauchline and Fisher (1969) review the earlier observations. The principal species occurring there are Thysanoessa raschi and T . inermis, although Meganyctiphanes norvegica is fairly common in its southern regions (Geiger et al., 1968; Degtyaryova et al., 1976; Kashkin, 1976; Drobysheva et al., 1977; Bliznichenko et al., 1978). Thysanoessa longicaudata penetrated the Barents Sea in comparatively large numbers in 1975 (Drobysheva et al., 1977). The changing abundance of Thysanoessa species has been examined over the period 1954-1972 by Drobysheva and Soboleva (1976) who conclude that their recruitment rate is dependent upon the environmental temperatures. Prygunkova (1974) records T . raschi in coastal regions of the White Sea. North Atlantic, north of about 40”N Thysanoessa raschi, T . inermis, Meganyctiphanes norvegica and Nyctiphunes couchi are common in the southern part of the Norwegian Deep (Fraser, 1970). Baan and Holthius (1969) monitored the plankton occurring around the lightship “Texel” at 53”N, 01’E in the southern North Sea between 1961 and 1966. Nyctiphunes couchi was the dominant species with Meganyctiphanes norvegica occurring occasionally. The former species was virtually absent from the surface plankton in the period May t o August, as it appears to be throughout the entire southern North Sea (Glover, 1952). They suggest t h a t this species is recruited from areas to the west and south-west of the British Isles by two routes, through the English Channel and from the north-west round northern Scotland. Fraser (1970) and Fraser and deaton (1969) describe the overflow of oceanic plankton t o the shelf waters of the north-east Atlantic. The species used as indicators of these events are primarily siphonophores but also the euphausiids Thysanopoda acutifrms, Euphuusia krohni, Nematoscelis megalops, Nematobrachion boiirpis, Stylocheiron longicorne, S. maximum and S. abbreviatum. Other recent work has been concerned with the euphausiid populations of the Gulf of St Lawrence (Eerkes, 1976, 1977a, b; Sameoto, 1976%)b), where the four commonest species are Meganyctiphanes norvegica, Thysanoessa inermis, T . longicaudata and T . raschi. Offshore, eastwards of

11. ECOLOGY

OF DISTRIBUTZON

517

Newfoundland and Labrador, the warm-water oceanic species Thysanopoda acutifrons, Euphausia krohni, Thysanoessa gregaria and Nematoscelis megalops occur along with the four St Lawrence species (Jones, 1969). Of these species only Thysano&sa inermie, T . longicaudata and T . raachi extend far up the Davis Straits. Mediterranean Sea Wiebe and D’Abramo (1972) examine the quantitative distribution of eleven species of euphausiids throughout the western and eastern basins of the Mediterranean. These basins are separated by Italy, Sicily and the shallow submarine ridge extending from Sicily t o Tunisia. Meganyctiphanes norvegica is largely restricted to the northern parts of the western basin as confirmed by Casanova (1970, 1974) and Bouchet and Thiriot (1972) who also found similar restriction on the distribution of Nyctiphanes couchi. The dominant species throughout the western basin were the temperate Euphausia krohni, Nematoscelis megalops and Stylocheiron longicorne ; Thysanoesa gregaria only occurred in this basin. The corresponding dominants in the eastern basin were the tropical-subtropical species Thysanopoda aeqwlis, Euphausia brevis and Stylocheiron suhmi. The euphausiid fauna of the Tyrrhenian Sea, the north-east segment of the western basin, is more closely related to the fauna of the eastern basin than t o that of the rest of the western basin. The euphausiid fauna in tihe eastern basin in the vicinity of the northern end of the Suez Canal is vary different from that of the northern Red Sea (Casanova et al., 1973). Sipos (1977) found that the dominant species of the eastern basin are not the dominant species in the Adriatic Sea. Meganyctiphunes norvegica, Nyctiphanes couchi, Euphausia krohni and Stylocheiron longicorne are dominant and these are the dominant species in regions of the western basin. Thysano&sa gregaria was not found in the Adriatic. The dominant species of the eastern basin are, with the exception of Ewphuusia brevis, sub-dominants in the Adriatic; El. brevis was not found there. Two other species, Stylocheiron abbreviatum and 8.maximum, occurred in the Adriatic in small numbers. Atlantic Ocean (4O”N-4O0S) Gopdakrishnan (1974) and Brinton (1975) include new records of the occurrences of various species throughout the Atlantic and these have been included where appropriate in Chapter 3. One of the most detailed analyses of an euphausiid assemblage is that produced by the “Discovery” SOND cruise of 1965 in the region

518

THE BIOLOGY OF EUPHAUSIZDS

of Fuertaventura, Canary Islands. Baker (1970) found 28 species, 20 of which were in sufficient abundance in the samples for Angel and Fasham (1973, 1974) to examine their occurrence in more detail statistically. The commonest species in this area are Euphausia krohni, E. hemigibba, Thysanohsa parva, Nematoscelis microps/atlantica, Stylocheiron longicorne and S. elongatum. Other common species are Thysanopoda aequalis, Euphausia brevis, E . gibboides, Nematoscelia tenella, N . megalops, Nematobrachion Jlexipes, Stylocheiron suhmi and S . afine. The great Meteor Seamount lies west of the Canaries a t approximately 30"N, 28'30'W. Weigmann (1974b) found 20 species of euphausiids in the region around the plateau of the seamount, the dominant species being Thysanopoda aequalis, Euphausia brevis, E . hemigibba, Stylocheiron suhmi and S. longicorne. Only 7 of these 20 species occurred on the plateau region, dominated by Euphausia brevis and Stylocheiron suhmi, the depth of water being considered insufficient to allow colonization by the other 13 species. The euphausiid fauna in the area south of the Canaries, off Cape Blanc, north-west Africa, has been studied by Weigmann-Haass (1976) and Andreu (1976). This is a region of coastal upwelling. The species composition of the fauna was similar to that of the Canaries and Meteor Seamount. Meira (1970) recorded 7 species, including the boreal Thysanohsa longicaudata, present among the Cape Verde Islands. Further to the south-east, in the Gulf of Guinea, Kinzer (1969b) mentions the occurrence of Euphausia tenera, E . eximia, E . mutica and Stylocheiron suhmi. Binet (1976), in discussing the distribution of species that he does not name, concludes that changes in the distributions of euphausiids across and along the shelf off the Ivory Coast are related ta variations in the Ivorian undercurrent. Individuals of a species are occasionally found well north of their distributional area in the North Atlantic; these are referred to as southerly guesks by Mauchline and Fisher (1969) who make little attempt to explain such occurrences. A related problem has been investigated in some detail by Wiebe and Boyd (1978) and Boyd et al. (1978). The southern boundary of occurrence of the cold slope species Nematoscelis megalops in the north-western Atlanbic is usually marked by the Gulf Stream. Individuals of this species, however, are recorded south of the Gulf Stream, in the Sargasso Sea. Such individuals were found ti0 be inside Gulf Stream cold core rings, associated with temperatures close to 10°C. As the rings age and warm so the euphausiids are forced to follow the cold temperatures downward6 to deeper environments. Their physiological condition deteriorates,

11.

ECOLOGY O F DISTRIBUTION

519

as indicated by the higher water and lower lipid contents of their bodies, and they finally die. In the western Atlantic, James (1970)recorded 30 species of euphausiids in the Gulf of Mexico, 21 of these being new records. Nine of these species were found in the Caribbean Sea by Owre and Foyo (1972) while Moore and Sander (1977), working in the extreme east of this region near Barbados, record 8 of the species. The species composition of these western Atlantic assemblages is closely similar to that of the Canaries-Cape Blanc region. A very different euphausiid assemblage is found on the Argentine shelf in the south-west Atlantic (Ramirez, 1971, 1973, 1977). The number of species is markedly reduced to about 10 and includes southern species such as E . lucens, E . vallentini, E . similis and Thysanoessa vicina.

North Pacijic, north of about 40"N There is little new information on the euphausiids of this region excepb the observations of Geiger et al. (1968) in the adjacent Arctic Basin, and Zhuravlev (1976b, 1977) and Shvetsova (1977) on the fauna of the Sea of Okhotsk. The species occurring in the Sea of Okhotsk are Eqhausia pacijica, ThzJsanohsalongipes, T . inermis and T . raschi. Shvetsova describes their distribution and biomass. Day (1971) describes the distribution and biomass of Euphausia pacijica, Tessarabrachion oculatum, Thysanoessa spinifera, T . longipes, T . raschi, Nematoscelis diflciicilis, Nematobrachion jlexipes and Stylocheiron maximum in bhe coastal region of the north-eastern Pacific extending from Vancouver Island southwards to Willapa Bay, Washington.

Pacijic Ocean (4O"N-4O0S) Maps of the distributions of adult and larval euphausiids in the region of the California Current have been supplied by Brinton (1967a, 1973) and Brinton and Wyllie (1976). Seasonal changes in the biomass of euphausiids throughout this region in 1955-1959 are given by Isaacs et al. (1969, 1971) and Fleminger et al. (1974). Variation in the biomass of euphausiids present was small and appeared to have no annual or seasonal pattiern. The fauna was dominated by copepods and salps, irrespective of season, these organisms representing 60-80 % of the wet biomass. Euphausiids represented 5 1 0 % of the total biomass, their mean biomass ranging from 1-15 g/lOOO m3. The distribution and abundance of some 30 species of adults are described

520

THE BIOLOGY O F EUPHAUSIIDS

and charts of the distributcions of the larvae of the dominant resident species, Euphausia pacijica, Nematoscelis dificilis and Thysanoessa gregaria and the dominant species with variable distributions throughout the region, Nyctiphanes simplex, Euphuusia gibboides, E . eximia, Thysanoessa spinifera and Stylbcheiron afine are given. The distributions of the different euphausiid species occurring along a transect between 23"N and 3"s in the Eastern Tropical Pacific (ETP) are described by Brinton (1979). Four distinct regions are present within this transect : ( 1 ) bhe California Current-ETP transition region off Baja California and the mouth of the Gulf of California; (2) the region between 22"N and l0"N which is characterized by oxygen deficient deep water; (3) the region of the North Equatorial Countercurrent ; (4) the equatorial region. Distinctive groups of species are associated with each of these regions. Brinton describes the horizontal and vertical distributions of the larvae and adults of each of the species and discusses the significance of these distributions in zoogeographical terms. The occurrence of euphausiids in the south-eastern Pacific in a band extending 150 miles seaward off the coast of Chile in latitudes 30"s to 42"s is described by Antezana (1970). The dominant species are Nyctiphanes simplex, Euphausia mucronata, E . vallentini and Thysanoessa gregaria. The spectrum of species, 14 in all, is similar to that found between 40"sand 50"s on the eastern coast of the South American Continent by Ramirez (1971, 1973, 1977). Roger has made detailed investigations in a series of papers of the tropical and southern tropical species in the region between 9O"W and 170"E and between the equator and 25"s. He defines faunal regions in the east-west and north-south axes of this sea area. The species composition of samples was different in the region of bhe equator from that in the tropical region between 15"s and 24"s. Further the species composition changed along the equator, the change arising from the presence of eastern, western and trans-Pacific species. The equator is richer in numbers of individuals than the tropical region; the density of euphausiids increases from west to east on the equator and east to west in the 15OS-25"S region. Further to the south, in Cook Strait, New Zealand, and the area to the south of it, Bartle (1976) and Robertson and Roberts (1978) found an euphausiid fauna consisting of 13 species of' diverse origin. The tropical and subtropical species, Stylocheiron elongutum, Euphccusia recurva and E . spinifera occurred along with the endemic Nyctiphanes australis and the southern species Euphausia lucens and E . vallentini. Sheard (1965, 1967) examined the complex region of the

11.

ECOLOGY OF DISTRIBUTION

621

eastern Australian Continental Shelf between 23"s and 44"s and describes the distributions of 12' species in this region in terms of the temperature and chlorinity characteristics of the water. Knox (1970) publishes data of J. A. Grieve describing the seasonal occurrence of several subtropical and sub-Antarctic species in the coastal waters of New Zealand. The species detailed are Nyctiphanes australis, Thysanoiksa gregaria and Nematoscelis species of warm-water origins and Euphausia uallentini from the sub-Antarctic. The dominant euphausiids occurring off Korea, that is in the western region of the Sea of Japan and the north-eastern region of the East China Sea, between latitudes 38"N and 33"N, are Euphuusia pacifica, E. nana and Pseudeuphausia latifrons; a further 7 species occw but in smaller numbers (Hong, 1969). This region has relatively few species and is very different from the South China Sea and Gulf of Thailand where Brinton (1975) and Rottman (1978) list 43 tropical and subtropical species. Many of these 43 species are derived from the adjacent tropical and subtropical Pacific and their presence in this region is a result of its oceanographic complexity. It is impossible t o summarize the contribution of Brinton (1975) to the zoogeography of euphausiids in an adequate manner. Much detailed and general information is given and the reader must refer to the original work.

Indian Ocean (toabout 40"s) Detailed distributional maps of the euphausiids of the Indian Ocean are now available (Gopalakrishnan and Brinton, 1969; Brinton and Gopalakrishnan, 1973 ; Brinton, 1975; Ponomareva, 1972, 1975). There are some significant differences between the distributional maps of some individual species as described by Ponomareva on the one hand and Gopalakrishnan and Brinton on the other ; these differences have been mentioned in the section on species distribution in Chapter 3. The different euphausiid assemblages tend to occur in latitudinal bands across the Indian Ocean. The region north of 10"N (Fig. 26), the Arabian Sea and Bay of Bengal, contains relatively few species dominated by the coastal Pseudeuphausia latifrons. Stylocheiron indicum, a coastal species with a restricted distribution, replaces X. longicorne in this area. The only other common species here are Euphausia diomedeae, Nematoscelis gracilis, Stylocheiron carinaturn and S. afline. Weigmann (1970) found the same species important in the Arabian Sea with the addition of Euphausia sibogae as defined by Briiiton (1975). Several species, namely Pseudeuphuusia latifrons,

522

THE BIOLOGY OF EUPHAUSIIDS

rr
-------

---__

____ - _ _ _

Boundary between subtropical and ternperole zones

- - - - - - - _

,

\

t

i

I

1

I

I

t

/

I

FIQ.26. Boundaries of euphausiid species in the Indian Ocean according to Brinton and Gopalekrishnan (1973).

Euphausia diomedeae, E . sibogae, E . sanzoi and 8.afine, occur in the Gulf of Oman but only one, Pseudeuphausia latifrons, appears to be present in the Persian Gulf (Weigmann, 1970, 1971). Weigmann (1970, 1974a) records P. latifrons, Euphausia diomedeae, E . sibogae, E. sanzoi, Stylocheiron a , n e and S. abbreviatum as present in the Red Sea, four of which occur as far north as the Gulf of Eilat. The 10"N latitude in the Indian Ocean forms the northern limit (Fig. 26) of some 8-10 tropical and subtropical circumglobal species listed by Brinton and Gopalakrishnan ( 1973). Euphuusia pseudogibha and Thysanopoda mtylata occupy the O0-l0"N band of latitude. The equator, which forms the southern edge of the North Equatorial Current, is the northern limit of Euphamia brevis and Thysanopoda aequalis. Latitude 10"s is the southern edge of the easterly Equatorial Counter Current, during November t o March, and, in May to September, the northern edge of the westerly South Equatorial Current

11.

ECOLOGY OF DISTRIBUTION

523

is at 6"-10"s. Species such as Euphausia diomedeae, E . paragibba, a markNemutoscelis gracilis and S t y Z o c ~ ~ r omn i c r o p ~ t ~ l mdecrease edly in numbers south of 10"s.Euphausia diomedeae is replaced by E. mutica and E . brevis ; Nematoscelis atlantica and S.fylocheironmicrophthalma are replaced by S. suhmi; and Euphausia paragibba is replaced by E . hemigibba. Species such as E . brevis, E . mutica, Stylocheiron suhmi and Thysanopoda aequalis have their southern limit of distribution between 25% and 30"s. This is also the southern limit of some other more widely ranging tropical and subtropical species and is the northern limit of Thysanoessa gregaria and the southern population of Euphausia similis. Finally, latitudes 40"s and 45% form the boundary region between subtropical and temperate species (Fig. 26). It is the southern limit of Euphausia recurva, E . hemigibba and Nematoscelis megalops. More detailed studies of particular regions of the Indian Ocean have been made. McWilliam (1977) examines the euphausiid assemblages along the 1lO"E meridian between latitudes 9'30's and 32"s. Euphausia brevis, E . mutica, E . tenera, Stylocheiron afine, S. carinatum and S. suhmi are the most abundant and ubiquitous species. She concludes that food resources rather than temperature and salinity influence tihe distributions, abundances and associations of species. Talbot (1974) identified 29 species in the Agulhas Current region off South Africa. She suggests that Thysanopoda tricuspidata, Stylocheiron suhmi and S. microphthalma are indicators of Agulhas Current water, while Nyctiphanes capensis and Euphausia lucens are indicators of waters of other origin. De Decker (1973) identified 34 species of euphausiids of the Agulhas Bank, west of the region studied by Talbot. Nyctiphanes capensis and Euphuusia recurva were the only species found on the Bank throughout the year, The latter species appears to replace the more offshore species, E . lucens, which only occurs over the Bank in winter months.

Antarctic Ocean Useful summaries of the general conditions in the Antarctic are given in Holdgate (1970). The general characteristics and distribution of Antarctic plankton are described by Voronina (1968) and Voronina and Naumov (1968). The standing stock is similar in all sectors and is dominated by copepods, the overall composition being similar to that in boreal regions. Voronina estimates that the meridional component of the current in the upper 100m layer moves the maxima of some species northwards from the Antarctic divergence at a mean

524

THE BIOLOGY OF EUPHAUSIIDS

velocity of 20 cmls. The water sinks at the Antarctic convergence at a rate of about 10.5 m/day, resulting in an approximate 3% increase per day in the total plankton in this region. Latogurskii et a2. (1975), working in the area of the Antarctic divergence between 52'E and 113'E, suggest that concentrations of krill are associated with the periphery of the zones of upwelling. Euphausia superba is now being fished as an economic resource and this has stimulated much work directed at defining its distribution and occurrence. Recent papers are listed for this species in Chapter 3 and a further discussion of its distribution is given in Chapter 13. Euphausiids aggregate but there are several types of aggregations. They can be relatively disorganized and be pdmarily caused by the physical features of the environment such as the presence of a front. They can be patches, shoals or surface swarms (sometimes called surface rafts). Patches are a normal feature of the distributions of many, if not most, planktonic organisms and may to some extent be generated by dynamic physical factors of the environment (Wiebe, 1960; Haury, 1976a, b). Shoals are large scale patches actively formed and maintained through behavioural mechanisms of the species ; they can be a seasonal feature-breeding aggregation-or a more permanent feature of the population. Swarms are usually smaller, more integrated patches of euphausiids with densities of animals measured in thousands per m3. This term is used here particularly in the context of surface and sub-surface swarms or rafts of species such as Euphausia superba. Swarms are occasionally stranded on shores and are recorded as strandings. Komaki (1967) reviews early observations of surface swarms and shore strandings and these have been combined with more recent observations in Table XXII. Mauchline and Fisher (1969) discuss the occurrences of these aggregations in some detail. Species such as Meganyctiphanes norvegica and Nyctiphanes australis form breeding aggregations prior to the breeding season but the latter species also occurs in swarms during the breeding season. Other species such as Thysanoksa gregaria are known to swarm a t the surface in large rafts during the breeding season but also a t other seasons. Baker (1970) observed a swarm of Euphausia krohni that may have been for purposes of breeding as did CasanovaSoulier (1970b) in the case of Meganyctiphanes norvegica. Euphausia pacijica frequently swarms in the northern regions of Japan (Komaki, 1967) but Brinton and Wyllie (1976) sbate that swarming is rare in this species in the California Current region. The greatest density recorded in the California Current was 20.4 per ms. This density is

TABLEXXII. AUUREGATION IN SPECIESOF EUPHAUSIIDS

Species

Number of animale Type of aggregation (per mS)

Thysanopodu tricuspidata shoal Meganyctiphanes norvegica swarms, strandings Nyctiphaw awrtralia swarms, breeding swarms N . capemis swarms Euph& eximia swarms E . krohni sub-surface shoal (220 m) shoal E . brevis E . diomedeae shoal, breeding swarms (S) E. rwuma patches, breedkg swarms swarms, sub-surface E . superba swarms E . vallentini Swarms shoal E . lucena E. j r i g i d a shoal swarms, shoals E . p,aci&u E . nana swarms swarms E . crystallorophiae E. aimilia swarms E. sibogae (distinguenda) E. spinifera T h y S a m h a spinifera swarms T . longipes swarms T . inapinata swarms T. immi.9

SW&rnS

T . tongicaudata T . gregaria T . raachi Nenmtoscelia megalops N . di@ilia Stylocheiron abbreuiatum

swarms, strandings swarms

0.02

5, 23 6, 9, 13, 24 6, 17, 21

31 000 20-100

19,22 23 12 18

7-15 0.8

2 23, 26

1, 16, 19, 26 c. 70 000

6, 7, 8, 11, 14, 16

75 000

11 19 2 6, 10, 25 10 3 10,19 5 19 6 6 9 6, 10, 20, 27 4, 20 6, 19, 26 6, 20 19 1, 25 23

150

600 000

SWBITnS

shoal shoal shoal

References

> 0.6 0.5

References: 1, Sheard, 1963; 2, Cushing and Richerdson, 1956; 3, Merr, 1962; 4, Fomfih and Jones, 1966; 5, Sebastian, 1966; 6, Komaki, 1967; 7, Ozawa et al., 1968; 8, Avfiov et al., 1969; 9, Mauchline and Fisher, 1969; 10, Nemoto et al., 1969; 11, Ragulin, 1969; 12, Baker, 1970; 13, Casanova-Soulier, 1970b; 14, Makarov et al.. 1970; 16, Wiebe, 1970; 16, Makarov and Shevtsov, 1971; 17, Bradford, 1972; 18, Boyd, 1973; 19, De Decker, 1973; 20, Zelickman and Golovkin, 1972; 21, Kawakami et al., 1973; 22, ,Cram and Schulein, 1974; 23, Roger, 1974c; 24, Cox, 1975; 25, Brinton and Wyllie, 1976; 26, Kawamure. 1977; 27, Kulka and Corey, 1978.

526

THE BIOLOGY OF EUF'HAUSIIDS

not indicative of a swarm unless the net is towed for an appreciable distance on either side of i t ; densities in swarms are usually measured in thousands per m3 (Table XXII).Swarms are probably formed in different species for different reasons and probably in some species only very occasionally. The formation of surface or sub-surface swarms has been reported in species that live predominantly in higher latitudes (Table XXII). Swarming has not previously been considered as an important featme of populations of tropical and subtropical species. Kawamura (1977), however, has found evidence of day-time swarming of Euphausia diomedeae, E . recurva and Thysanoessa gregaria in the south-western Indian Ocean (30°S), the Coral Sea (lo's),and in the south-western Pacific Ocean (25'5) where they were consumed by Bryde's Whale. The seasonal and sexual composition of the swarms suggests that the Euphausia species may form breeding aggregations. Nyctiphanes capensis forms surface swarms off the south-western African coast at about 22'5 at night. Cram and Schulein (1974) estimated the area of the swarms to be 86.1 km2. The euphausiids were luminescing and restricted to the upper 36m of the water column. This is the largest recorded surface patch or swarm of an euphausiid; the densities of the animals within the swarm were not determined. Further information is required on the occurrence of aggregation within populations of euphausiids. Observations that should be made are identification of the species involved, the area and thickness of the aggregation and the density of animals within them. Surface swarms of Euphausia superba are of especial interest in the context of commercial exploitation of this species. Observations of swarming of this species are discussed in Chapter 13 and some of %hecharacteristics are probably pertinent to swarms of other species.

CHAPTER 12

PREDATORS A N D PARASITES The predators of euphausiids are numerous and Mauchline and Fisher (1969) have enumerated them in considerable detail. There have been, however, several recenti investigations of considerable interest. The discussion of the predators follows in the same order as that in the previous work, namely, whales, seals, fish, squid, decapod crustaceans, other invertebrates and birds. Nemoto (1970a) has produced an account of the comparative structure of the baleen plates related to diet in the baleen whales. He finds three feeding types: swallowers, skimmers and those skimming and swallowing. The swallowers (blue, fin, Bryde's, humpback, minke whales) have cavernous mouths with internal volumes as great as 4-6 m3. They fill the mouth with water and euphausiids and then discharge the water through the baleen plates while retaining the euphausiids which are then swallowed. This feeding behaviour was filmed by Gaskin (1976) who describes it graphically in a fin whale off south-western Nova Scotia. They never swim with their mouths open, a featme of the skimmers whose feeding is a more continuous process. The skimmers (right, Greenland and pygmy right whales) sieve a continuous stream of water that enters the open mouth and passes through the baleen plates, the euphausiids or other organisms being retained. These whales do not require swarms of organisms but can feed on much more sparsely distributed food. The sei and grey whales appear to use a combination of these two feeding methods. The various species of baleen whales range over wide geographical regions and consequently show considerable variations in their diets (Gaskin, 1976). The following euphausiids are important constituents of the diet of whales in the North Pacific: Euphausia pacijca, Thysanoiksa inermis, T . longipes, T . spinifera and T . raschi. South of 40'N in the northern Pacific, Euphuusia pacijca remains important but E . similis, E. recurva and E . nana replace the more northern Thysanoiksa species in the diets (Nemoto, 1970a). The fin and sei whales north of 40"N in the Pacific fed on Euphausia pacijca and south of 40"N the diet of the sei changed t o include E. recurva, E. diomedeae,

528

THE BIOLOGY OF EUPHAUSIIDS

E . tenera and Nematoscelis dificilis; these whales had a mixed diet of copepods, euphausiids, decapods and fish (Kawamura, 1973). Antezana (1970) states that Euphausia mucronata and Nyctiphanes simplex are important food organisms of the fin whales off the coast of Chile. Rorvik et al. (1976) suggest that the. distribution of fin whales in the North Atlantic is closely related to that of Meganyctiphanes norvegica. Kawamura (1977) determined the food of Bryde's whale in the warm regions south of Madagascar ( 3 0 ° S ) , in the Coral Sea (10's) and in an area north of New Zealand (25'5). This whale inhabits regions with surface water temperatures of approximately 20°C and feeds exclusively on Euphausia diomedeae, E. recurva and Thysanokksa gregaria. Other recent work contributes further information on the food of whales in the Antarctic. Zenkovich (1970) and Mackintosh (1970) estimate that before commercial exploitation the whales consumed about 150 million tons of planktonic crustaceans, mainly euphausiids, per season. Schott's Southpolar regions of the three oceans, as defined in Fairbridge (1966), correspond reasonably well with the region under discussion. Assuming the total 160 million tons to be euphausiids and that they occur in the upper 200m layer, this crop is equivalent to about one euphausiid per 10 m3 over the whole region. The standing stock of euphausiids is probably closer to one individual per m3 and so the original whale seock would be cropping about 10% of the euphausiid stock per season. Euphausia superba is the most important euphausiid in the Antarctic. Brown (1968) reports that the sei whales ate this species exclusively around South Georgia. The sei whales in the region of Crozet and Kerguelen Islands usually feed exclusively, according t o Kawamura (1970), on one of the following organisms: Calanus tonsus, Parathemisto gaudichaudi, Euphausia superba or E. vallentini. The food of this whale in the Indian and western Pacific sectors of the sub-antarctic can vary although Kawamura (1974a) found that more than 70% fed exclusively on one species of prey organism. The most important euphausiids in the diet were E . valbntini, E . lucens and E. superba but some E . diomedeae and Thysanqoda acutifrons were also present. The small minke whales in the Antarctic were found to be feeding on Euphawia superba and E . spinifera by Ohsumi et al. (1970). Only one size group usually occurred in any one stomach. Mackintosh (1974)' examining the stomach contents of blue, fin, sei and humpback whales, suggests that the whales tend to select swarms of krill containing larger individuals in a year class. The most heavily predated size class of euphausiids were in the body length range 2760 mm.

12.

PREDATORS AND PARASITES

629

Mauchline and Fisher (1969) have listed the euphausiid8 eaten by whales in different geographicd regions. The following species must now be added t o this list: Euphausia nana, E . recurva, E. diomedeae, E . tenera and Nematoscelis dificilis in the north Pacific; Nyctiphanes simplex and Euphausia mucronata off Che coast of Chile; E . lucens, E . diomedeae and Thysanopoda acutifrons in the subantarctic. Thysanobsa vicina is also eaten in the Antarctic (Kawamura, 1974s). This means that 27 species have now been recorded as included in the diets of whales. The crabeater and leopard seals of the Antarctic are listed by Oritsland (1970) as feeding on euphausiids. Sergeant (1973) found that the north-west Atlantic harp seal, Pagophilus groenlandicus (Erxleben), feeds on Meganyctiphanes norvegica, capelin and herring in the Gulf of St Lawrence and on Thysanobsa inermis and T . raschi and other foods in the Arctic. A great variety of fish feed on euphausiids. Mauchline and Fisher (1969) discuss predation by herring, Clupea harengus L., haddock, Melanogrammus aeglejinus (L), saithe, Pollachius virens (L), and other gadoids including the cod, Gadus morhua L. Nagabhushanam (1965) summarizes the information on the diets of some Irish Sea and North Atlantic gadoids, noting the importance of species such as Meganyctiphanes norvegica. Euphausiids are one of the main sources of nutrition for cod in the Barents Sea in summer, the species consumed being ThysanoGsa raschi, T . inermis and T . longicaudata (Soboleva, 1975) ; euphausiids comprised 6 6 5 % of the stomach contents of 0-group fish (Ponomarenko, 1976, 1978). Meganyctiphanes norvegica is a component of the diets of gadoids in Passamaquoddy Bay (Tyler, 1972). A deep benthic gadoid, the longfin hake Phycis chesteri Goode and Bean, eats Bentheuphausia amblyops, Meganyctiphanes norvegica and Stylocheiron elongatum according to Sedberg and Musick (1978). Macpherson ( I97 8a, b) recorded ~ e g a n y c t i ~ h a nnorvegica, e~ Eu~hausia krohni, Nyctiphanes cowhi and Nematoscelis megabps as components of the diets of Phycis blennoides (Briinich), Antonogadus megalokynodon (Kolombatovic), Gadiculus argenteus argenteus Guichenot and Micromesistius poutassou (Risso) in the western Mediterranean. Another gadoid, Micromesistius australis, eats large quantities of Euphausia superba in the Scotia Sea (Permitin, 1970). Pitt (1973) found Meganyctiphanes norvegica and Thysanobsa species to be a minor component of the diet of the plaice, Hippoglossoides platessoides (Fabricius), on the Grand Banks, Newfoundland. The Pacific sanddab, Citharichthys sordidus (Girard), the rex sole, Glyptocephalus zachirus Lockhgton, and the slender sole, Lyopsetta exilis (Jordan and Gilbert),

530

THE BIOLOGY OF EUPHAUSIIDS

feed on Euphausia pacijka off the Oregon coast (Kravitz et al., 1976; Pearcy and Hancock, 1978; Vanderploeg, 1979), as does the arrowtooth flounder, Atheresthes stomias (Jordan and Gilbert), according t o Vanderploeg (1979). The halosaurid Halosauropsis macrochir (Giinther) eats Thysanopoda species and Bentheuphausia amblyops, while the synaphobranchid Synqhobranchus kaupi Johnson eats Thysampoda acutifrons, T . microphthalma, Stylocheiron elongatum and S . maximum on the continental slope and rise of the Atlantic coast of the United States (Sedberry and Musick, 1978). Several skates are known to feed to some extent on euphausiids. Meganyctiphanes occurs among the stomach contents of Raja erinacea Mitchill, R. ocellata Mitchill, R . radiata Donovan and R . senta Garman, in the western Atlantic coastal region between Nova Scotia and Cape Hatteras (Tyler, 1972; McEachran et al., 1976). Holden and Tucker (1974) found that euphausiids are an important item in the diets of young R. clavuta L. and R . montagui Fowler and a minor item in the diet of R. brachyura Lafont in British waters. Barracouta, tuna and albacore eat euphausiids. The barraoouta, Thyrsites atun (Euphrasen), feed on Nyctiphanes simplex off the coast of Chile (Antezana, 1970) and on N . australis in the eastern Cook Strait, New Zealand (Mehl, 1969). The striped tuna, Katsuwonw pelamis (L), eats Nyctiphanes simplex off the coast of Chile (Antezana) while the slender tuna, Allothunnus fallai Serventy, feeds on Nyctiphanes australis in Tasmanian seas (Wolfe and Webb, 1975; Bishop et al., 1976); N . australis comprised 96% of the stomach contents of the slender tuna. Roger and Grandperrin (1976) examined the stomach contents of 365 albacore, Thunnus (Germo) alalunga (Gmelin), and yellowfin tuna, T . (Neothunnus) albacares (Bonnaterre), in the tropical Pacific; they found some 1400 micronektonic fish which had been feeding to a considerable extent on a number of species of euphausiids (see below). Yellowh, skipjack (striped) and other tuna feed to a small extent on euphausiids in the temperate and tropical Atlantic (Dragovich, 1969, 1971 ; Matthews et al., 1977). Information on the importance of micronektonic fish as predators of euphausiids is now available. Roger and Grandperrin (1976) examined some 1400 fish, the species being listed by Legand et al. (1972), obtained from the stomachs of tuna (see above). The stomachs of half these fish contained recognizable stomach contents ; euphausiids of the genera Stylocheiron, Euphausia, Thysanopoda and Nematoscelis comprised half the diet of 25% of these fish. The principal predators on euphausiids among these epipelagic fish are in the families Gempylidae, Paralepididae and Bramidae (Roger, 1973e). The species of

12. PREDATORS

AND PARASITES

531

euphausiids eaten were : Thysanopoda tricuspidata, T . monacantha, Euphausia fallax, E . diomedeae, 2. tenera, Nematoscelis microps, N . gracilis, N . tenella, Stylocheiron abbreviatum, S . maximum, S . longicorne, S . mrinatum, S . suhmi, S . elongatum, S . afine, and S . microphthalma. Studies have been made of the food of the deeper-living micro. nektonic fish, the dominant families of which are the Myctophidae, Gonostomatidae and Sternoptychidae. Roger (1 973d) examined the stomach contents of species in these three families and also in the Chauliodidae, Bregmacerotidae and Nemichthyidae in the tropical Pacific. None of these fish feeds exclusively on euphausiids but Roger estimates that the species listed above in the genera Thysanopoda, Euphawia, Nematoscelis and Stylocheiron comprise about 10% of the diet. Clarke (1978) records euphausiids among the stomach contents of the myctophids Benthosema suborbitale (Gilbert), Ceratoscopelus warmingi (Lutken), Lampanyctus niger (Gunther), L. nobilis, L. steinbecki and Triphoturus nigrescens, the gonostomatid Gonostoma atlanticum Norman and the photichthyid Vinciguerria nimbaria (Jordan and Williams) in Hawaiian waters. Merrett and Roe (1974) investigated the food of Argyropelecus h e m ~ g y ~ n u Cocco, s A . aculeatus Cuvier and Valenciennes, Chauliodus danae Regan and Trewavas, Valenciennellus tripunctulatus (Esmark), Lobianchia doJleini (Zugrnayer), Noto1ychnu.s valdiviae (Brauer) and Lampanyctus cuprarius Taning, in tihe eastern North Atlantic. Copepods were the dominant item of the diet but euphausiids were eaten by all species. Thysanopoda obtusifrom,T. aequalis, Nemutoscelis microps and Stylocheiron Eongicorne were identified in the stomachs of Chauliodus danae. Hopkins and Baird (1977) analysed the diets of 17 species of myctophids, gonostomatids and sternoptychids from various regions of the Atlantic and also from the Pacific and Indian Oceans. They discuss ontogenetic, seasonal and regional variation in the diets. Euphausiids were only identified t o genus and occurred commonly in many of the fish. Samyshev and Schetinkin (1973) found that the myctophids Diaphus dumerili (Bleeker), D. taaningi Norman, Lepidophanes guentheri (Goode and Bean) and the gonostomatid Maurolicus muelleri (Gmelin) fed on euphausiids, frequently as a major portion of the diet, along with copepods and decapod larvae off north-west Africa. A detailed study of the diet of the myctophid Benthosema glaciale (Reinhardt) in western Norway demonstrated that it feeds on euphausiids, especially Thysanoha species and 0-group Meganyctiphunes norvegica, throughout the year (Gjosaeter, 1973). The food of this myctophid off north-western Africa consisted principally of copepods and ostracods but the larger fish,

532

THE BIOLOGY OF EUPHAUSIIDS

over 30 mm in length, fed on Euphausia krohni in the surface layers at night (Kinzer, 1977). The food of myctophids in the Pacific off Oregon (Tyler and Pearcy, 1975) and California (Paxton, 1967) included E . pacijica, Thysanoessa spinifera and Nematoscelis dificilis. Permitin (1970) found Euphausia superba in the stomachs of about 26% of the Antarctic myctophids Protomyctophum tenisoni and Electrona antarctica (Giinther) and as the dominant food of a larger species Gymnoscopelus nicholsi (Gilbert). There is evidence that more temperate species such as Paralepis atlantica prionosa and Anotopterus pharao Zugmayer perform feeding migrations to feed on Euphausia superba in Dhe Antarctic. A detailed stiudy of the diet of the hatchet fish, Sternoptyx diaphana Herman, by Hopkins and Baird (1973) showed that euphausiids of the genera Euphausia, Thysanoksa, Nematoscelis and Stylocheiron were dominant items in the Atlantic and subantarctic Pacific Oceans. Many other fish prey on euphausiids (Mauchline and Fisher, 1969). Haedrich and Henderson (1974) and Sedberry and Musick (1978) found evidence of the deep benthic rattail, Nematonurus armatus (Hector), feeding pelagically on Meganyctiphanes norvegica, Nematoscelis megalops, N . microps and Bentheuphausia amblyops. Studies off Oregon by Pearcy and Ambler (1974) show that large macrourids feed mainly on pelagic organisms, among them Thysanopoda species. They review the literature on feeding of macrourids and note that euphausiids occur fairly frequently; this was confirmed by McLellan (1977), who lists 22 macrourids with euphausiids in their diets. The osmerid eulachon, Thaleichthys pacijicus (Richardson), feeds on euphausiids off Oregon (Vanderploeg, 1979). The blue ling, Molwa byrkelange Walbaum, consumes euphausiids in the north-east Atlantic (Koch and Lambert, 1976). Trachurus japonicus (Teraminck and Schlegel), the jack mackerel, eats Euphusiapacijica off Japan (Suzuki, 1973). The Pacific saury, Cololabis saira (Brevoort), consumes several species (Fukushima, 1976; Odate, 1977), Merluccius gayi gayi (Guichenot) eats Euphausia mucronata off the coast of Chile (Antezana, 1970), M . capensis Cast. feeds on euphausiids off South Africa (Bentz, 1976) and Cram and Schulein (1974) record surface shoaling of this species to feed on swarm8 of Nyctiphanes cqensis. Taius tumifrons (Temminck and Schlegel) feeds on euphausiids in the Yellow and East China Seas (Enomoto, 1972) and Arripis trutta preys on swarms of Nyctiphanes australis a t Kaikoura, New Zealand (Bradford, 1972). Parsons and Le Brasseur (1970) state that juvenile pink salmon, Oncorhynchus gorbuscha (Walbaum), consume small quantities of Euphausia pacijca off western Canada, while Andrievskaia (1974) found that Thysanoksa Eongipes along with

12.

PREDATORS AND PARASITES

533

amphipods is its main diet in the winter in the Sea of Japan. Okada and Tmiguchi (1971) state that the chum salmon, Oncorhynchus keta (Walbaum), feeds on a mixed diet of organisms that includes Euphausia p"ci$ca, Thysanoessa longipes, T . inspinata and T . raschi off northern Japan. Inshore fish do not normally have access to many euphausiids. Species such as Neganyctiphanes norvegica, Euphausia pacifica, Thysanoessa inermis and T . raschi do, however, occur in many coastal regions and are consumed by local populations of fish. Hobson and Chess (1976) state that Thysanoessa spinifera move into shallow water at nighti when they are eaten, as a relatively minor component of mixed diets, by the olive rockfish, Sebastes serranoides (Eigenmann and Eigenmann), the kelp rockfish, S. atrovirem (Jordan and Gilbert), the queenfish, Seriphzrs politus Apes, and the blacksmith, Chrornis punctipinnis (Cooper). Two South African coastal fish feed to a small extent on Euphausia recurva and E . lucens; these are the hottentot, Pachymetopon blochii (Valenciennes), and the silverfish, Argyrozona argyrozona (Valenciennes), studied by Nepgen (1977). Mauchline and Fisher (1969) briefly discuss predation of krill by Antarctic species of fish. Permitin (1970), Naumov and Permitin (1973) and Permitin and Tarverdieva (1978) provide further information on the feeding of notothenids and chaenichthids on Euphausia superba and suggest that not only Antarctic fish but several subantarctic and subtropical fish, by making feeding migrations, also feed on E . superba. Shust (1969) observed Notothenia rossi murmorata (Fischer) jumping among surface swarms of Euphausia superba on which i t was feeding. Latogurskii (1972) states that krill are the main component of the diet of this fish but Tarverdiyeva (1972) found it to feed t o a larger extent on ctenophores. Two Antarctic percoid fish, Trematomus borchgrevinki Boulenger and T . newnesi Boulenger, feed on krill and other crustaceans under the ice (Andriashev, 1970). Raja georgiana Norman eats Euphausia superba off South Georgia (Permitin, 1970). Legand et al. (1972) review information on the feeding of squid. Kawakami et al. (1973) state that Nototodurus sloani sloani (Gray) feeds on swarms of Nyctiphanes australis and the amphipod Parathemist0 gaudichaudi off New Zealand. Another squid, Todarodes pacificus (Steenstrup), feeds on Euphausia pacijica in the northern Pacific (Kawamura, 1974b). The pelagic decapods have been found to feed on euphausiids. Meganyctiphanes norvegica, Euphausia krohni and Nyctiphanes couchi off the Moroccan Coast in the north-east Atlantic are one of the main constituents of the diets of the following decapods (Lagardere, 1972,

534

THE BIOLOQY OF EUPHAUSIIDS

1977a, b) : Pmiphaea sivado (Risso), P . multidentata Esmark, Plesionika kterocarpus (Costa), P. martia (A. Milne-Edwards), Chlorotocus crmsicornis (Costa), Processa canaliculata Leach, Philocheras echinulatus (M. Sars), Aristeus antennatus (Risso), Aristeomrpha foliacea (Risso), Plesiopenaeus edwardsianus (Johnson), Solenocera membranacea (Risso), Parapenaeus longirostris (Lucas), Polycheles typhlops Heller, Nephrops norvegicus (L.), Munida sarsi Lagardere (1973, 1977b) found that Dicklopandalus bonnieri Caullery in the Gulf of Gascogne fed to a notable extent in May and August, and predominantly in July, on euphausiids; no euphausiids occurred among their stomach contents in March and April when they were feeding on fish. The most common species of euphausiid was Meganyctiphunes norvegica but Nyctiphunes couchi and Euphuusia krohni were also recognized in the diet. Donaldson (1975) states that Sergestes splendens Sund, 8. atlanticus H. Milne-Edwards, S. corniculum Kroyer and S. grandis Sund prey to some extent on euphausiids near Bermuda; several other sergestids did not eat euphausiids. Another selection of decapods was examined by Foxton and Roe (1974) in the region of 30"N, 23"W, west of the Canary Islands. Species of euphausiids, which were unidentified, occurred among the stomach contents especially of the larger individuals of the following species: Acanthephyra purpurea (A. Milne-Edwards), Systellaspis debilis (A. Milne-Edwards), Gennadm valens (Smith), Sergestes atlanticus H. Milne-Edwards, S. henseni (Ortmann), ' S . grandis Sund, S. 'pectinatus 'Sund and S . sargassi Ortmann. Cannibalism may be practised t o a small extent within some populations a t certain times. Adults of a species may occasionally eat their furciliae, or adults or furciliae of another species, as indicated by the presence of parts of euphausiid eyes within their stomach contents (Table VI, p. 450). Pavlov (1976a) records the presence of parts of the bodies of Euphuusia superba among the stomach contents of E . triacanthu in winter months when other food was scarce. Deep sea copepods may prey to some extent upon euphausiids. Harding (1974) found that the guts of Pseudochirella polyspina Brodsky, 1950 and Scaphocalanus magnus (T. Scott, 1894) contained large portions of the eyes of euphausiids. Crystalline cones were present in the hindguts suggesting that the predation was real and that the copepods had not just simply attacked the euphausiids in the plankton bucket during the course of capture. Moguilevsky and Gooday ( 1977) record euphausiids occasionally among the stomach contents of the crustacean ostracod Gigantocypris muelleri Skogsberg ; they suggest that these are genuine occurrences

12. PREDATORS AND PARASITES

535

and that they were not eaten in the plankton buckets. The species of euphausiids concerned were hot identified. The lobate ctenophores Ocyropsis crystallina (Rang, 1828) and 0. mculata (Rang, 1828) have been observed eating euphausiids in the laboratory (Harbison et al., 1978).

Mauchline and Fisher (1969) review information on predabion of euphausiids by birds. Zelickman and Golovkin (1972) found that the little auk Plotus alle L. feeds on Thysanohsa longicaudata, T . inermis and T . raschi off the northern shores of Novaya Zemlya. The least, crested and parakeet auklets-Aethia pusilla (Pallas), A. cristatella, (Pallas) and Cyclorrhynchus psittacula (Pallas)-feed on surface swarms of adults and young Thysanoessa inermis and T . rmchi during the summer in the area of St Lawrence Island, Alaska (Bedard, 1969). Analyses of sea bird stomachs in New Zealand by Bartle (1976) showed Nyctiphanes australis to be an imporCant food for the grey-faced petrel Pterodroma macroptera (Smith) and the fairy prions Pachyptila turtur (Kuhl). Larus novaehollandiae scopulinus Stephenson also fed on this euphausiid (Bradford, 1972). Gudkov (1962) listed birds associated with patches of euphausiids and other plankton in the Barents Sea. The guillemots, Uria species, feed on a mixed diet that includes Thysanohsa longipes and T . rmchi in the eastern Bering Sea (Ogi and Tsujita, 1973). A list of Antarctic birds congregating on patches of Euphausia superba is given by Ozawa et al. (1968); 17 species are listed. Shust (1969) found that the Pintado petrel, Daption capensis L., was the commonest bird associated with patches of Euphausia superba. Other birds present were the Antarctic petrel, Thalmsoica antarctica (Gmelin) the wandering albatross, Diomedea exulans L., the black-browed albatross, D. melanophris Temminck, the Antarctic skua, Catharacta antarctica (Lesson), the giant petrel, Macronectes giganteus (Gemelin) and Wilson’s storm petrel, Oceanites oceanicw (Kuhl). The information on the predators of the different species of euphausiids given in Mauchline and Fisher (1969) and the present paper is summarized in Table XXIII. Discussion of the importance of predation on certain species of euphausiids is given in the former work. References to predation on euphausiids are scattered and difficult to find. Many papers refer to euphausiids in general and provide few details of the species involved. Some useful conclusions, however, can be drawn from Table XXIII. All species of euphausiids probably aggregate to some extent at certain seasons, times or in certain regions. Such aggregations are more readily detected in coastal and surface swarming species and

TABLEXXIII.

PREDATORS OF

EUPHAUSIIDS Fish

Species

Aggregating Whales Seals

BentheuphaMia arnblyopa Thysanopoda mnacantha T . criatata T . tricwpidata X T.aeqUa2ia T. aatylata T . obtuaifrons T . pectinatu T . orientalis T . rnierophthalm T . acutij+rons T . cornuta T . egregia T . spinicaudata X Meganyctiphanes norvegica Nyctiphanee couchi ? N . auatralis X N . capen& X N . simplex ? Paeudeuphaausia lcctifrons ? P. &nim Euphataia americam E. exirnia X E . krohni X E. mutica X E. brevis E. diornedeae X E . recurva X E. Buperba X E. vallentini X E. lucens X E . frigida X E. pacifica X E. nana X E. crystdlorophiaa X E. tenera X E. similis E. similia var. arnata E. mucronata E. sibogae X E . distinguenda E. larnelligera E. gibba

D

P

M

DecaSquid pods Birds

+

+ +

+ +

+

+ + +

+ + + +

+ + + + t+ + + + + +

+ + + + + + + +

+ + + + + + + + + + +

+

+

+

+

+

+

+

+ +

+

+

+ + + + + + +

f

+

+ + +

+

+

+

+

+ + + + + + contd.

TABLEX X I I I - e o n t d . Fish Species E. gibboides E. fallax E. sanzoi E. pseudogibba E. paragibba E. hemigibba E. spinifera E. hanseni E. longirostris E. trimantha Teasarabrachion oculatum Thysanoessa spinifera T . longipes T . inapinata T . inermis T . longicuudata T . parva T.gregaria T . wicina T . rnmwra T . raschi Nematoscelt dificilis N . megalops N . tenella N . microps N . atlantim N . lobatu N . grmilis Nematobrachion pexipea N . sexspinosum N . boopis Stylochiron carinaturn S. afine S. suhmi S . microphthalma S . insulare S. elongatum S . indicum S. longicorne S . abbreuiatum S. maximum S. robwtum

Aggregating WhalesSeaL

D

P

DecaSquid pods Birds

M

+

+

+ x

+

+ + + + + + + + +

X

+ + + + + +

+ + + + + +

+

+ + +

+ + + +

+

+ + +

+ +

+ + + +

+ + + + + + + + + + + + + + + + + + + + + +

Species listed in Table XXII as aggregating are shown ( x ); those not listed in Table XXII but which probably aggregate are indicated (?) ;fish are divided into demersel (D), planktivorous (P) and micronektonic (M).

538

THE BIOLOGY OF EUPHAUSIIDS

less easily recorded in sub-surface oceanic populations. Aggregation is known to occur in 19 of the 27 species listed as food for whales. Four of the other 8 species not listed as aggregating are Nyctiphanes simplex, Pseudeuphausia latifrons, Thysanobsa vicina and T . macrura. The f i s t two are coastal species and so probably do aggregate in common with all other coastal species. Likewise, aggregation is a general feature of Thysanobsa species (Table XXII) and so these two Antarctic species probably behave in the same way. The other four species of non-aggregators are Thysanopoda acutifrons, Euphausia tenera, E . mucronata and E . hemigibba. The first species has only been recorded once in a whale stomach; E . tenera is eaten simultaneously with other euphausiids in the north Pacific, E . mucronata is eaten by whales in the coastal region of Chile where it may aggregate locally and E . hemigibba was found in the stomach contents of a humpback whale along with other species. Consequently, whales are feeding only on species that form distinct aggregations. It is these species in the genera Meganyctiphunes, Nyctiphanes, Pseudeuphausia, Euphuusia and Thysanoessa that are also most heavily predated by seals, demeraal fish, planktivorous fish, squid and, of course, sea birds. The food of pelagic decapods has only been investigated in north-eastern Atlantic and Pacific species. Four species of euphausiids have been identified in their stomachs (Table XXIII)but further studies will undoubtedly increase this list, probably to include Thysanopoda and Nematoscelis species and some of the deeper-living Stylocheiron species. The micronektonic fish are listed (Table XXIII) as preying on 16 of 34 species in the genera Thysanopoda, Nematoscelis and Stylocheiron and only on 10 of the 51 species in the other 7 genera. The former 3 genera contain predominantly oceanic, cosmopolitan species many of which live in the meso- and bathypelagic environments (Table IV, p. 431). These species are not available as food for demersal fish or indeed for most populations of planktivorous fish such a8 sardine, herring, mackerel, etc. Some tuna feed on euphausiids through consuming micronektonic fish. Most of these euphausiids do not form aggregations in the surface layers and so are not eaten by whales or birds. Pelagic decapods probably do-eat them but their most important predators are probably the micronektonic fish. Marshall (1971) suggests that the ratio of numbers of euphausiids t o micronektonic fish in the mesopelagic environment is probably in the range 3-5 t o 1. There may be some potential competition for food between these relatively small fish and the euphausiids. Roger (1973~)points out that the position of the euphausiids in the food webs will depend upon their abundance and accessibility to predators. Accessibiliby is

12. PREDATORS AND PARASITES

539

probably primarily controlled by the bathymetric distribution of the species in combination with their body size. Only two parasites of euphausiids are described in Mauchline and Fisher (1969), the first a crustacean bopyrid Branchiophryxus nyctiphanae (Caullery) on Meganyctiphanes norvegica and the ellobiopsid Thalassomyces f q e i Boschma on several species of euphausiids. Much further information on euphrtusiid parasites is now available. There are three types of ectoparasites of euphausiids : ellobiopsids, dajid isopods and apostome ciliates. The occurrence of the first two types is shown in Table XXIV. The development of Thalassomyces f q e i has been described (Mauchline and Fisher, 1969); this parasite does not necessarily prevent development of the secondary sexual characteristics although it does tend to castrate its host. These parasites have been classed as fungi, dinoflagellates or rhizopod protozoans (Mauchline and Fisher, 1969). Recently, Galt and Whisler (1970) noted that T . marsupii, parasitic on the hyperiid amphipod Parathemisto pacijicu Stabbing, produced biflagellate spores. They concluded that the number and constant orientation of the flagella and the general morphology of the spores suggest a relationship with the achlorophyllous Dinophyceae. Field ( 1969) has studied the biology of Notophryxus lateralis which parasitizes Nematoscelis ditficilis in the Santa Barbara Channel, California. It causes castration of both males and females. Field suggwts that the parasite inhibits moulting in the euphausiids and a similar inhibition must be caused by Thalassomyces fqei. The phoronts, or resting cysts, of apostome ciliates have been recorded attached to Meganyctiphanes norvegica, Nyctiphanes cowhi, Euphausia hemigibba, E . krohni, Thysanoifssa gregaria, T . inerrnis, T . longicaudata, T . raschi and Nematoscelis megalqs by Lindley (1978b) in the North Atlantic. Infection ranged from 3% to 16% of the population. Lindley describes the geographical distribution of infested euphausiids and concludes that neritic species and populations are more liable to infection than offshore oceanic species and populations. There are several recorded endoparasites of euphausiids (Table XXV). Komaki (1970) has reviewed the earlier literature. Nematodes ascribed to Anisakis species occur in euphausiids, which are the intermediate hosts, and in whales, which are the final hosts. Fish feeding on euphausiids become infected with Anisakis larvae (Wooten, 1978). Oshima (1969) experimentally infected Euphuusia pacijim and E . similis with Anisakis larvae. Shimazu and Oshima (1972) demonstrated natural infection by Anisakis larvae in various species of

540

THE BIOLOGY OF EUPHAUSIIDS

TABLEXXIV. ECTOPARASITES OF EUPHAUSIIDS Parasite

Euphausiid

References

Ellobiopsid

Thalassomyces fagei (Boschma)

Dajid Isopods Branchiophryxua nyctiphanae (Caullery) Heterophqxua appendiculatua G. 0. Sars Notophryxua globularis G. 0. Sars N . lateralis G. 0. Sars

Branchiophryxus koehleri Nierstrasz & Brender B . caulleryi Koehler

Meganyctiphanes norvegica Nyctiphanes auatralis N . capensis Euphawia krohni E . diomedeae E. recurva E . vallentini E . lucens E. frigida E . pacifim E . tenera E. similis E. sibogae E . sanzoi E . pseudogibba E . hemigibba ThysanoZssa longipes T . inermis T . gregaria T . rmchi Nemtoscelis dificilis

6, 9, 11 6 10,12 6 8 6 6 6 6 13 8 6 8

Meganyctiphanes norvegica

4

8

6

5, 6 13 14 6 13 6

Euphausia krohni E. sibogae Thysanoessa gregaria Nemtoscelis dificilis N . megdops Stylocheiron earinaturn 8.afine S. longicorne

References: 1, Sam, 1885; 2, Koehler, 1911; 3, Field, 1969; 4, Mauchline 8nd Fisher, 1969; 5, Baker, 1970; 6, Komaki, 1970; 7, Sebastian, 1970; 8, W e i p a n n , 1970; 9, Franqueville, 1971; 10, Meira, 1971; 11, De Bhaldraithe, 1973; 12, Vader, 1973; 13, Wing, 1976; 14, Lindley, 1977.

euphausiids in the North Pacific, Uspenskaya (1963) having previously recognized infection of ThysanoGssa rasehi in the Barents Sea. Smith (1971) and Lindley (1977) found infected ThysanoZssa species in the North Sea but Kagei et al. (1978) found no Anisakis larvae in two examinations of large samples of Euphausia superba consisting of

12.

541

PREDATORS AXD PARASITES

TABLEXXV. ENDOPARASITES OF EUPHAUSIIDS PWasite Nematoda (larvae) Anieakk sp.

Ascurophis sp. Contracaecum sp. (Type B ) Contracaecum sp. (Type D ) Trematoda fistomurn Jilijerum G. 0. Sam

metacercaria metacercaria o f P a r o n d r e w sp. Cestoda (plerocercoids) Nybelinia aumnenicola Okada

Euphazcsiid

Thysanolisea longicccudata T. inemis T . raachi T. longipea Euphawia paci,fica Thyaanohaa i n e m i s T. raachi Euphauaia p i J i c a E. p a c i m ThysanoZasa raachi

Rejerencee

4 4, 5, 7 2, Ei 5 5

7 5

5 5 5

ThysanoEasa gregaria Newtoscelia megalops EuphauSia Sirnilis E . paci$ca Euphawia pacifica T h y s a n o h a longipea T . inemis T . raachi T . longipea

Pelkhnibothrium catdatum Zschokke and Heitz Cystocercoide T. inemis Acanthocephala (larvae) Echinorhynchue cormcgatzcs G. 0. Sara E u p b w i a krohni Bolbosoma c a e n o j o m (Heitz) Thysanohsa longipes ’ T . raechi Palaeoacanthocaphatlan T . longicawhta

6

References: 1, Sam, 1885; 2, Uspenkaye, 1963; 3, Komaki, 1970; 4, Smith, 1971; 5, Shimazu end Oshima, 1972; 6, Shimazu, 1975; 7, Lindley, 1977; 8, Shimazu and Kagei, 1978.

36,319 and 55,295 individuals respectively. Komaki (1970) has been the first t o recognize the presence of trematode metacercariee in an euphausiid; Sard records of infection are from the South Atlantic, Komaki’s from the North Pacific. The records of cestode and acanthocephalan larvae tire principally from the North Pacific but that of Lindley is from the north-east Atlantic; the location of Sam’ record is unknown.

CHAPTER 13

EUPHAUSIIDS IN THE MARINE ECONOMY Mauchline and Fisher (1969) have listed the species of euphausiids that are important in tihe marine economy of the various regions of the oceans. This list is still valid as a list of the most important species. Additional species are named in Table XXIII of this review; many of the new ones are very prominent in the diets of the populations of cosmopolitan micronektonic fish, il sitiuation that has only been realized in the last few years The species of euphausiids involved are the oceanic members of the genera Thysanopoda, Tessarabruchion, Nematoscelis and Stylocheiron One other predominantly oceanic genus, Nematobrachion, has not been recognized as food for these fish; this is surprising because Ohe three species in the genus are frequently common in samples Measurements of biomass or standing stocks of euphausiids are reviewed in Mauchline and Fisher (1969). The primary conclusion was that euphausiids represented a major fraction of the totial biomass of plankton, this biomass being proportionally higher in higher latitudes where the eight or nine most important species are resident. More recent measurements of the biomass of euphausiids have been made in widely diverse sea areas. The densities found range from 0.1 to 10 individuals per m3. The euphausiids usually constitute 5-10% of the total biomass of plankton and about 30% of the biomass of the crustacean plankton. No conclusions of any real value, additional to those given in Mauchline and Fisher (1969), can be drawn at this stage and so the relevant papers presenting biomass measurements are simply listed according to the Ocean in which the observatiions were made: Atlantic: Greze et al. (1969); Egan and Conrad (1975); Sameoto (1976b); Berkes (19774 ; Blackburn (1977); Lindley (1978a). Puci$c : Nakamura (1967); Shomura and Nakamura (1969) ;Brinton (1967a, 1973); Isaacs et al. (1969, 1971); Blackburn et al. (1970); Vinogradov (1970); Legand et al. (1972); Roger (1973c, 1978a); Vinogradov and Parin (1973); Fleminger et al. (1974); Maynard et al. (1975); Brinton and Wyllie (1976); Murano et al. (1976); Blackburn (1977); Brinton (1979).

13.

EUPHAUSIIDS IN THE MARINE ECONOMY

543

Indian : Gopalakrisban and Brinton (1969).

Antarctic: Korabel’nikov (1975). The present data on euphausiids, especially Euphusia pacijica, are nearly sufficienb to examine the economics of a population. The model of growth, described by Mauchline (1976, 1977th) and used in Chapter 10, forms the starting point. Mauchline (1977b) calculated curves for various parameters of an individual E . pacijica against time (Fig. 27). The curves relating the decay of the percentage growth factors, based on measurements of body length and dry weight, are

Time (days1

FIU. 27. Euphuwia paci$m. Curves of selected growth parameters throughout the life of the animal. Length factor shows the decline in the percentage increase in body length at successive moults and weight factor shows the corresponding decline in the percentage increase in wet weight. Growth rate shows the actual daily increment in body dry weight; broken line indicates the period over which the ovary would develop if the daily growth increment remained at 0.07 mg/day. The curves for increase in body length and dry weight are given along with a curve of the weight of the casts produced with time. The curve for weight specific oxygen consumption is superimposed. The durations and timing of the successive intermoult periods are shown. The ~ F P O W Sdenote the position of the relevant scales (after Mauchline, 1977b).

544

THE BIOLOGY O F EUPHAUSTIDS

shown along with the durations af the successive intermoult periods. The growth curves relating the increase in body length and dry weight have been generated in the manner shown in Table XX (p. 506) for E . superba. The weights of successive casts can be plotted and the curve for weight specific respiration rate superimposed. The growth increment, calculated as mg tissue per day, can be plotted and shows that it reaches a maximum value and then declines. The timing, within the life history of the animal, of the achievement of maximum increments coincides with the development of the ovary which, when mature, represents approximately 10% of the body weight. The rate of growth of the ovary has been determined in several species (Mauchline and Fisher, 1969); some 8-16 weeks is required for full development, the period being dependent upon the species and its environmental temperature. Approximate daily increments in weight for the ovarian tissue can be calculated such that a mature ovary will be produced within the required period. These increments are always approximately equal to the maximum increments in body weight calculated in the growth curve. Storage products, such as lipids, are often utilized during the period of ovarian development (Mauchline and Fisher, 1969). The relative contributions of food intake and storage products to the development of the ovary have not been determined. Further, no experimental information on the weights of food eaten per unit time diurnally and seasonally is available. The weight specific food intake curve, weight specific respiration curve, and the weight specific excretion curve are probably of the same form. There can, of course, be “perturbations” within the animal’s physiology that will affect these curves, together and separately. Examples of such perturbations, all on different scales of time, are the generation of storage or reproductive products, horizontal migration and diurnal vertical migration. The effects of perturbations in the animal’s physicalchemical environment on its physiology also have different time scales, ranging from sporadic short term variations to recurrent seasonal cycles. Allowances and correction factors for such perturbations, once recognized and quantified, can be made in models of the animal’s physiology. Assimilation efficiencies have been measured in a few instances in euphausiids (see Chapter 9) and range from 65% to %yo, dependent to some degree on the type of food. Much more information is required, especially in relation to assimilation of the variety of seasonal diets of the animal in the sea. Gross growth efficiency, K,, defined as the percentage of captured food that is converted into new tissue and net growth efficiency, K,, defined as the percentage of assimilated food that is converted into new tissue, have not been determined. The factors affecting these parameters also require

13.

545

EUPHAUSIIDS IN THE MARINE ECONOMY

investigation. Further, more data are also required on rates of excretion. It is possible, however, to use curves such as t,hose in Fig. 27 to estimate parameters of populations (Mauchline, 1977b). The data on the individual in Fig. 27 can then be integrated to provide measurements of the population as a whole, for instance a P : B ratio. The usefulness of the concept of P : B ratio is discussed by Mauchline (1977b). This is the ratio of annual production (P) to mean annual biomass ( B ) . The P : B ratios of several populations of euphausiids have been calculated (Table XXVI). This ratio is primarily dependent upon the size structure of the population and the duration of the season of active growth. Individuals in the population of Euphausia pacijica off Oregon are growing and reproducing a t all seasons of the TABLEXXVI. P : B RATIOSIN EUPHAUSIID POPULATIONS Speciea Euphausia pacifica

E. superba Meganyctiphanes norvegica ThysanoBsa raschi T . longicaudata

Sea area

Oregon California Antarctic Scotland St. Lawrence N. Atlantic

P

:B

ratio

8.7 3 1.8-2.3 2.3 4 1.2-11.6

Reference Mauchline (1977b) Mullin (1969) Allen (1971) Mauchline (197713) Berkes (1977a) Lindley (1978a)

year. The species is relatively small in body size and any single individual does not live longer than one year. This is reflected in a high P : B ratio of 8.7. Thysanoessa longicaudata in the northern Atlantic has P : B ratios ranging from 2 : 1 to 3 : 1 in areas where a single generation is produced per year. Ratios, however, of 2.4 : 1 to as high as 11.6 : 1 occur in regions where %wogenerations are produced annually (Lindley, 1978a). Mullin (1969)made an estimate of the ratio in Euphausia pacijica off California but his value of 3.0 is too low. Allen’s value (Table XXVI) of 1.8-2.3 for the Antarctic species E. superba is in accordance with those for Meganyctiphunes norvegica and Thysanoessa raschi a t lower latitudes. These species all have life-spans of up to two or more years and live in environments dominated by marked seasonal fluctuations. Mauchline and Fisher (1969) discuss the Antarctic food chains in relation to the transfer of vitamin A, because of the crucial role of euphausiids, especially Euphausia superba, in this region. One of the most interesting and recent investigations of food chains is that

546

THE BIOLOGY OF EUPHATJSIIDS

in the tropical Pacific by Legand et al. (1972) and Roger (1968-1975). This investigation has shown that euphausiids are important in the diets of the micronektonic fish which in turn are fed on by the tunas. The euphausiids represented S-lO% of the diet of the fish. The structure of the food chains was determined not only from the vertical distributions and migrations but also from the feeding rhythms of the animals. Roger and Grandperrin (1976) attempt the construction of a functional model of this tropical food web (Fig. 28). The model is simplified by being discretely divided into day and night. The feeding rhybhms of the animals are not as regular, as has been shown 0

200

400

4 8

rn

8

8

v 8

4

4

60C 4

8

8

8

T

8 4

q

80C

0phytoplankton;

,o non-migrant zooplankton (e.g. Sly/acheifon spp.); a ,o non-migrant long-line tunas and their food, the micronektonic fish; 8 non-migrant deep fauna ;A,A vertically migrating zooplankton (e.g. Euphusiu and rhySQnupGdu spp. 1; , a , 8 , A feeding; 0 , 0 ,A non-feeding. FIG.28. The pelagic assemblage of the tropical Pacific showing (arrows) the main paths of energy transfer (after Roger and Grandperrin, 1976).

13.

EUPHAUSIIDS IN THE MARINE ECONOMY

547

for euphausiids by Roger in Fig. 19 (p. 455). The model does, however, illustrate the primary parameters in the vertical plane. The migrating fauna feeds on the epipelagic fauna a t night and Roger and Grandperrin argue that since the epipelagic fauna does not, also feed primarily at night there is the equivalent of an “energy valve”. It encourages the downward transfer of energy and reduces any upward transfer. This is an interesting hypothesis. It,appears to assume the presence of excess food for the epipelagic fauna so that they have no requirement for more or less continuous feeding. Inactivity a t night may aid them to avoid predation by the migrant faunas. Roger and Grandperrin restrict this scheme to the tropical Pacific and, by inference, to other tropical regions. The epipelagic fauna could probably be expected to be more voracious in middle and higher latitudes where tihe food is seasonally scarcer. The possible role of faecal pellets of euphausiids in the downward transport of nutrients is briefly discussed by Mauchline and Fisher (1969). Smayda (1969) and Fowler and Small (1972) have examined the sinking rates of pellets. Natural faecal pellets, as opposed to those produced in the laboratory when the animals are fed on laboratory diets, sank at faster rates than the sinking rates of eggs and at comparable rates to those of cast integuments. The sinking rate increased with increasing volume of the pellets and ranged from 126 m/day to 862m/day; the range in volume of the individual pellets was 0.210 x l o 7 pm3. Fowler and Small (1972) conclude that euphausiid pellets may be at least as important as casts and carcasses in the downward transfer of nutritive matter to the deep layers and the benthic populations. Faecal pellets are probably extremely important in transferring concentrations of substances, such as PCB compounds, zinc, cerium, polonium, plutonium and lead isotopes, downwards in the water column, because these substances are concentrated in or on them (Elder and Fowler, 1977; Higgo et al., 1977; Beasley et al., 1978; Cherry et al., 1978). La Rosa (1976) has recently described a simple method of recovering faecal pellets in experimental systems. The possible contribution that euphausiids make to the marine economy through moulting and the consequent production of casts is discussed by Mauchline and Fisher (1969)on pages 383-385. Sameoto (1976b) studied the euphausiids of the Gulf of St Lawrence where densities as high as 1180 animals per m2 occur. He calculated that the quantity of casts produced was 25.9/mg/m2/24h or 77 cal/m2/24h. Ogawa (1977) discusses the role of faecal bacteria in not only degrading the pellets but also as a source of nutrition for the euphausiids.

548

THE BIOLOGY OF EUPHAUSIIDS

One aspect of the pelagic assemblages that has emerged is the complementary relationship be6ween mysids and euphausiids. Mysids are predominantly coastal organisms. They inhabit estuaries, lagoons and beach areas of up t o 30-50 m depth and are frequently pelagic in behaviour in these shallow areas. They become epibenthic or hyperplanktonic in regions with depths between 6Om and about 400m, the pelagic environment above them being inhabited by coastal and slope species of euphausiids. Some strictly pelagic species of mysids occur in this region but their densities in the water column are much lower than those of the euphausiids. I n oceanic regions, the euphausiids occupy the upper regions of the water column. The mysids, as far as is known, are not necessarily associated with the sea bed in abyssal depths but occupy a pelagic zone below the main distributional range of the euphausiids (Figs 29 and 30). There are, of course, a few epipelagic oceanic species of mysids and a few bathypelagic species of euphausiids. % plankton biomass

I/

% plankton biomass

10 0

.. .

.. .. .. _.

B

B

.:.

8t A

FIU.29. Euphausiids and mpids as a percentage of the total biomass of plankton st different depths. Left: A, euphausiids;B, mysids; solid lines, Kuril-Kamchatkeares; hatched lines, tropical Pacific. Right: A, euphausiids and B, mpids in the tropical Pacific (after Vinogradov. 1970; Vinogradov snd Parin, 1978).

13.

EUPHILUSIIDS IN THE MARINE ECONOMY

Of”

649

wet weight

FIG.30. Percentage vertical distribution of the biomess of left, euphausiids, right, myaids by day ( u n w e d ) and night (shaded) (after Murmo et al 1976).

The possibilities of exploiting the Antarctic Ewphausia euperba and other species are discussed in some detail by the following authors : Mauchline and Fisher, 1969; Makwov, 1970; Gulland, 1970; Mackintosh, 1970; Moiseev, 1970; Bogdmov, 1971; Le Gall and L’Heroux, 1972; Postel, 1972; Fischer, 1974; Bykov, 1975; Nemoto and Nasu, 1975; Sarhage and Steinberg, 1975; Pinhorn, 1976; Anonymous, 1977; Eddie, 1977; Everson, 1977; Freytag, 1977; Grantham, 1977; Roschke and Schreiber, 1977; Omori, 1978; Sarhage et al., 1978. Ewphausia p a c i j h is caught commercially in several regions, especially off northern Japan where it is dried and marketed for bait and feed for farmed fish (Fulfon, 1976; Mason, 1976; Koops et al., 1977). A bibliography on krill as a human food resource is provided by Grantham (1976, 1977) and Taylor (1976). There has been a large decrease in the numbers of baleen whales living in the Antarctic through overfishing. This has led to the belief that there is now present a significant excess crop of euphausiids that is no longer utilized by the whales. Estimates of the possible annual yield of a commercial fishery for E. superba range from 30 to 200 million tons (Gulland, 1970; Mackintosh, 1970; Allen, 1971; Anonymous, 1977; Everson, 1977; Kock and Neudecker, 1977; El-Sayed, 1978; El-Sayed and McWhinnie, 1979). Prisent opinion guesses that the yield is probably closer to

650

“HE BIOLOGY OF EUPHAUSIIDS

the upper than to the lower end of this range. Everson (1977) and El-Sayed and McWhinnie (1979) review the results of exploratory commercial fishing for E . superba and provide some catch statistics. Various methods of location and capture of surface and subsurface swarms of euphausiids have been proposed or tested (Groisman et al., 1969; Tupolev, 1969; Postel, 1972; Cram and Schulein, 1974; Koyama et al., 1974; Eddie, 1977; Everson, 1977; Freytag, 1977; Herring and Locket, 1978). They include echo-sounder and visual location of swarms, capture by ring or purse nets operable from small boats, large stern trawls and large trawls fixed to the side of ships with a pump mechanism drawing the euphausiids on board. The use of light lures of differing colours and intensities has been investigated. Semenov (1969) suggests that light can be used t o increase the density of euphausiids in a swarm rather than as a lure. The intensity of the light must be low because high intensities cause the swarm to disperse (Mikhailowkii, 1969). Petushko (1969) found that a red light attracted the krill whereas the normal coloured artificial light caused them to scatter and aggregate at the edge of the lit area in the twilight zone. Krill were found to be very sensitive to d.c. electric fields but experiments using an electric field adjacent to the side of the ship produced no swarming (Petushko, 1969). Once &he euphausiids are captured, they can be processed in several ways to yield large amounts of animal protein (Yanase, 1971, 1974a, b ; Arai e t - a l . , 1976; Watanabe et al., 1976; Grantham, 1977; Kuwano and Mitamura, 1977). The chemical composition of the species has already been described and methods of treatment and preparation are discussed by Lyubimova et al. (1973), Flechtenmacher et al. (1976), Roschke (1977) and Tokunaga et al. (1977). The krill contain large amounts of trace elements, vitamin A, “B” vitamins and importanb fatty acids (see Chapter 7). Krill pastes and meal (Lyubimova et al., 1973; Bulycheva et al., 1977; Fedotova et al., 1977; Leinemann and Christians, 1977; Jahn et al., 1978) can be manufactured and used as animal feed and in the therapeutic diets of patients suffering from stomach ulcers and arteriosclerosis. Lyubimova et al. (1973) and Grantham (1977) discuss the production of various manufactured products such as krill sausages, stuffed eggs and shrimp butter and suggest that there are many other potentially marketable recipes. The euphausiids have to be present in swarms to be commercially exploitable. It is only when swarming that sufficient weight per unit fishing effort can be obtained to make the operation financially profitable. Such a fishery depends upon the almost continuous location and capture of the swarms and it is thus of considerable importance to

13.

EUPHAUSIIDS IN THE MARINE ECONOMY

55 1

know the distributions of swarming populations in time and space. Avilov et al. (1969) found that swarms occur in association with fronts and also near underwater mountains and Bogdanov et al. (1969), Bogdanov and Solyanik (1970) and Makarov (1 972) describe concentrations of Euphuusia superba associated with the frontal zone in the Scotia Sea. The krill swarms were present when the biomass of phytoplankton was within the range 1.0-5.0 ml/m3 but were absent in areas where less than 0.5ml/m3 and more than 5*0ml/m3were present (Avilov et al., 1969). Shust (1969), however, concludes that the formation of swarms is independent of phytoplankton biomass; the persistence of the swarms, on the other h'and, in a sea area does seem to be dependent upon the presence of phytoplankton and the euphausiids continue swarming at the surface longer during daylight hours in regions with higher biomasses of phytoplankton. Pavlov (1969, 1974) suggests that swarms are formed by individuals which have fed to the full and have ceased feeding and that the duration of swarming is linked with the availability of food. Obviously, much further investigation of the environmental, and possibly physiological, parameters governing initiation of swarming of species such as E . superba and E . pacijica are required. Surface swarms of these two species are usually 5-100m in diameter (Mackintosh, 1966; Avilov et al., 1969; Nemoto et al., 1969; Ragulin, 1969). The thickness of swarms studied by Ragulin was 1 - 6 5 m and the krill were all orientated in the same diredion. Shust (1969) found very large surface aggregations of E . superba that he termed fields. They could be 1500-5500m in length and seemed to consist of aggregations of individual swarms that could be discerned within the field. Surface swarms of E . superba reacted to a diver a t a distance of 1.5-2 m but did not disperse. Sub-surface swarms occur most commonly at depths between 5 m and 5 0 m and the deeper layers under the swarms are often inhabited by salps (Avilov et al., 1969). There is substantial evidence that swarms me cohesive throughout 24 h and perform, as a unit, a diurnal vertical migration, occurring in the surface 20 m layer at night but moving downwards t o the 60100m layer during the day (Sasaki et al., 1968; Shust, 1969; Mohr, 1976). Pavlov (1974) on the other hand, observed swarms rising to the surface at midnight and a second time a t noon to feed ;they swarmed actively when not feeding but dispersed to feed. Shevtsov and Makarov (1969) and Mohr and Fischer (1977) found several patterns of behaviour. Consequently, swarms may behave irregularly. Surface swarms of krill can be seen visually at distances as great 8s 15002000m from a ship in dull weather but are usually absent during

552

THE moLoaY OF EUPHAUSIIDS

sunny weather (Shust, 1969): The presence of swarms is often advertized by congregations of sea birds feeding on them. The distance between individuals in a swarm is approximately equal to the cube root of the volume occupied by a single animal (Clubter, 1969)) assuming that the animals are evenly dispersed throughout the volume of the swarm. Everson (1977) has reviewed the information on bhe densities a t which E. superba occurs in swarms. The density range is 2000-60,000 animals/m3 of water and the corresponding range of distance between individuals would then be 2.5-8.0 em. Kils (1979) found that E . superba respires at a rate of approximately 1 pl O,/mg dry weight/h and that the critical environmental oxygen concentration is at about 90% saturation. Consequently, he points out that oxygen depletion within swarms will control their density, maintenance, size and movement. The density of phytoplankton will also influence the formation and maintenance of swarms (Mezykowski and Rakusa-Suszczewski, 1979). Mauchline and Fisher (1969) discuss the age composition of swarms of krill. Any one swarm is often composed of a relatively restricted size range of individuals. Shevtsov and Makarov ( 1 969) and Makarov (1970) found a clear spatial separation geographically of young and mature sbages of E . superba in the Scotia and northern part of the Weddell Seas. Makarov (1972, 1973, 1974b) and Makarov and Maslennikov (1975) have described the distribution of larvae and breeding adults and the dispersal patterns away from spawning areas in the Scotia Sea. A detailed analysis of the sizes of krill occurring in whale stomachs demonstrated seasonal, regional and annual variations in the sizes of krill available to the whales (Mackintosh, 1974). Whales consumed krill primarily of body length between 27 mm and 50 mm and imposed the heaviest mortality in the size range 15-40 mm. The geographical distribution of E . superba is described in very general terms in Chapter 3 and in Mauchline and Fisher (1969). It has been reviewed in some detail by Shevtsov and Makarov (1969)) Mackintosh (1973)) Voronina (1974) and Everson (1976). The details of its distribution, the recruitment and maintenance of populations in the Weddell Sea and East Wind Drift, annual and seasonal variations in its distribution are all areas where much further work is required. This is a reflection of the strong commercial interest in this species. It has also been realized that there is a lack of information on its general biology. The length of time required for it to achieve sexual maturity as well as its longevity are disputed. Ivanov (1970) suggested that it does not mature sexually until it is three years old,

13. EUPHAUSIIDS IN THE MARINE

ECONOMY

553

the majoriby of females breeding a t an age of four years; a small portion of trhe population may even reach an age of five years. Mackintosh (1972), on the other hand, concluded that it matures sexually and breeds at an age of two years; he does not comment on longevity but his analysis suggests that a small portion of the population may attain an age of three years and breed a second time. These very different conclusions were drawn by Ivanov and Mackintosh from essentially the same data, Mackintosh having the added advantage of examining Ivanov's paper before submitting his own for publication. This situation reflects the difficulties present in studying and interpreting datia on oceanic species. Such difficulties can usually only be resolved if samples can be taken repetitively from the same recognizable population in which all individuals have had the same environmental hist,ory. The krill species of the northern hemisphere, in the northern Atlantic and Pacific Oceans, are Meganyctiphanes norvegica (M. Sars), Euphausia pacijca Hansen, Thysano&sa inermis (Krrayer) and T . raschi (M. Sars). There are essentially isolated populations of all these species and it has been possible to study their dynamics and, through statistical analysis of samples of the populations, to provide good estimates of mean growth rates and longevity of individuals (Table XXI, p. 511). The only known isolated population of Euphawsia superba is that reported by Antezana et aE. (1976a) to exist in a fjord in southern Chile at approximately 50"s. No observations on the biology of this population have been published. Ivanov (1970) thought that each female E. superba bred during only one season of her life span but Makarov (1975b, 1976) has now demonstrated t.hat breeding can take place in ati least two successive seasons. Consequently, this species is similar to the North Atlantic species that are known t o breed in two successive seasons. An empirical analysis of the growth of E . superba is presented earlier in this review (Fig. 25, p. 509). There are seasonal, regional and annual variations in the rates of growth of the North Atlantic and Pacific species of euphausiids and such variations are undoubtedly characteristic of this Antarctic species. The effects of commercial exploitation on populations of an organism with a life cycle of two years will be different from the effects on populations with a cycle of three to four years. One of the primary reasons for this is that recruitment rates will be different. The shorter life cycle requires higher recruitment rates per unit time to maintain the population bhan in the case of the longer cycle. This directly affects the crop that can be harvested by commercial exploitation without reducing the stocks below non-viable levels.

REFERENCES Ackman, R. G., Eaton, C. A., Sipos, J. C., Hooper, S. N. and Castell, J. D. (1970). Lipids and fatty acids of two species of North Atlantic krill (Meganyctiphanes norvegica and Thysanoessa inemis) and their role in the aquatic food web. Journal of the Fisheries Research Board of Canada, 27, 513-533. Allen, G. H. (1972). Notes on sampling amphipods and euphausiids in the North Pacific Ocean. Transactions of the American Fisheries Society, 101, 577-582. Allen, K. R. (1971). Relation between production and biomass. Journal of the Fisheries Research Board of Canada, 28, 1573-1581. Alton, M. S. and Blackburn, C. J. (1972). Die1 changes in the vertical distribution of the euphausiids, ThysanoZssa epinif era Holmes and Euphausia pacifica Hansen in coastal waters of Washington. Calqornia Fish and Game, 58, 179-190.

Andersen, N. R. and Zahuranec, B. J., eds (1977). “Oceanic Sound Scattering Prediction”, Marine Science, Vol. 5, 859 pp. Plenum Press, New York and London. h d r e u , P. (1976). Contribucih a1 estudio de 10s eufausihceos de 10s alrededores de Cabo Blanco (NW de Africa). Resultados Expdiciones Cientiflcas del Buque Oceanografico “Cornide de Smvedra”, No. 5, 77-1 11. Andriashev, A. P. (1970). Cryopelagic fishes of the Arctic and Antarctic and their significance in Polar ecosystems. I n “Antarctic Ecology” (Ed. M. W. Holdgate), Vol. 1, pp. 297-304. Academic Press, London and New York. Andrievskaia, L. D. (1974). The feeding of pink salmon in the wintering areas in the Sea of Japan. Izvestiya Tikhookeanekogo Nauchno-issledovatl’skogo Instituta Rybnogo i Okeanografii, 90, 97-1 10. Angel, M. V. (1977). Windows into a sea of confusion: sampling limitations to the measurement of ecological parameters in oceanic mid-water environments. I n “Oceanic Sound Scattering Prediction” (Eds N. R. Andersen and B. J. Zahuranec), Marine Science, Vol. 5, pp. 217-248. Plenum Press, New York and London. Angel, M. V. and Fasham, M. J. R. (1973). Sond cruise 1965: factor and cluster analyses of the plankton results, a general summary. Journal of the Marine Biological Association of the United Kingdom, 53, 185-231. h g e l , M. V. and Fasham, M. J. R. (1974). Sond cruise 1965: further factor analyses of the plankton data. Journal of the Marine Biological Association of the United Kingdom, 54, 879-984. h o n y m o u s (1968). Monographs on Oceanographic Methodology. 2. Zooplankton sampling, 174 pp. United Nations Educational Scientific and Cultural Organization, Paris. Anonymous (1977). Biological investigations of marine Antarctic systems and stocks (BIOMASS), Krill. Scientific Committee on Oceanic Reaearch (Working Group 54), 24-34. h t e z a n a , T. J. (1970). Eufausidos de la costa de Chile. Su rol en la economia del mar. Revista de biologia marina, Valparaiso, 14, 19-27.

REFERENCES

555

Antezana, T. J. and Fowler, S. W. (1972). Gamma emitters in euphausiids from the southeast Pacific Ocean. Health Physics, 22, 201-205. Antezana, T., Aguirre, N. and Bustamante, R. (1976a). Nuevo limite septentrional para el krill Antarctico Euphausia superba Dana, en el Pacific0 (Crustacea, zooplancton). Series Cientifico Instituto Antarctico Chileno, 4, 47-51. Antezana, T., Aguirre, N. and Bustamante, R. (1976b). Clave ilustrada y distribucion latitudinal de 10s eufausidos del Oceano Antarctico (Crustacea, zooplancton). Series Cienti8co Instituto Antarctico Chileno, 4, 53-68. Antonini-Kane, J., Fowler, S. W., Heyraud, M., KefkeS, S., Small, L. F. and Veglia, A. (1972). Accumulation and loss of selected radionuclides by Meganyctiphanes norvegica M. Sars. Rapport et proc8s-verbaux des rdunions. Commission internationale pour l’exploration scientifique de la mer Mdditerrande, 21, 289-290. Arai, K., Watanabe, T. and Kinumaki, T. (1976). Studies on the utilization of Antarctic krill, Euphausia superba Dana-I. Nutritive value by a rat-feeding trial. Bulletin of the Tokai Regional Fkheries Research Laboratory,No. 85,l-12. Arbault, S., Beaudouin, J. and Lacroix, N. (1972). Zones-test dans le Golfe de Gascogne en 1970. Ichthyoplancton-zooplancton.Annalea Biologiques, 27, (1970), 71-72. Aron, W. and Collard, S. (1969). A study of the influence of net speed on catch. Limnology and Oceanography, 14, 242-249. Artiges, J. M., Pagano, M. and Thiriot, A. (1978). Morphologie fonctionnelle des appendices nutritionnels de Meganyctiphanes nowegica (M. Sars, 1856) et Euphaueia krohnii (Brandt, 1851) (Crustacea Euphausiacea). Archives de Zoologie Exphimentale et Ge’ndrale, 119, 95-106. Avilov, I. K., Elizarov, A. A., Kanaeva, I. P. and Lavrov, G. N. (1969). Biooceanological premises of the search for krill. Trudy Vseaoyuznogo Nauchnoissledovatel’skogo Imtituta Morskogo Rybnogo Khozyaktva i OkeanograJi, 66, 246-248. National Lending Library for Science and Technology, Boston Spa, Yorkshire, RTS 5691. Ayah, F. J., Valentine, J. W. and Zumwalt, G. S. (1974). Genetic variability in Antarctic krill. Antarctic Journal, 9, 300-301. Ayah, F. J., Valentine, J. W. and Zumwalt, G. S. (1975).An electrophoretic study of the Antarctic zooplankter Euphawia superba. Limnology and Oceanography, 20, 635-640. Baan, S. M. van der and Holthuis, L. B. (1969). On the occurrence of Euphausiacea in the surface plankton near the lightship “Texel” in the southern North Sea. Netherlands Journal of Sea Research, 4, 364-371. Bacescu, M. (1975). Contributions to the knowledge of the mysids (Crustacea) from the Tanzanian waters. Science Journal, University of Dar es Salaam, 1, 39-61. Bacescu, M. and Mayer, R. (1961). MalacostracBs (Mysidacea, Euphausiacea, Decapoda, Stomatopoda) du plancton diurne de la MBditerranBe, Btude bas& sur le material du Lamont Geological Observatory, Washington. Rapport et proc8s-verbaux des r6unions. Commission internationale pour l’exploration scientifique de la mer Mkditemande, 16, 183-202. Bainbridge, V. and Forsyth, D. C. T. (1972). An ecological survey of a Scottish herring fishery. Part V: the plankton of the north-western North Sea in relation to the physical environment and the distribution of herring. Bulletin of Marine Ecology, 8, 21-52.

556

THE BIOLOGY OF EUPHAUSIIDS

Baker, A. de C. (1959). Distribution and life history of Euphaueia trimantha Holt and Tattersall. Discovery Reports, 29,309-340. Baker, A. de C. (1970). The vertical distribution of euphausiids near Fuertaventura, Canary Islands (“Discovery” Sond Cruise, 1965). Journal of the Marine Biological Association of the United Kingdom, 50, 301-342. Baker, A. de C., Clarke, M. R. and Harris, M. J. (1973). The N.I.O. combination net (RMT 1 8) and further developments of rectangular midwater trawls. Journal of the Marine Biological Association of the United Kingdom, 53,

+

167-184.

Bamstedt, U. (1976). Studies on the deep-water pelagic community of Korsfjorden, western Norway. Changes in the size and biochemical composition of Meganyctiphanes norvegica (Euphausiacea) in relation to its life cycle. Saraia, 61, 15-30.

Bartle, J. A. (1976). Euphausiids of the Cook Strait: a transitional fauna? New Zealand Journal of Marine and Freshwater Research, 10, 559-576. Bary, B. McK. (1956). Notes on ecology, systematics and development of some Mysidacea and Euphausiacea (Crustacea) from New Zealand. Paei,jic Science, 10, 431-467.

Bary, B. McK. and Pieper, R. E. (1971). Sonic-scattering studies in Saanich Inlet, British Columbia :a preliminary report. In “Biological Sound Scattering in the Ocean” (Ed G. B. Farquhar), pp. 601-609. Maury Center for Ocean Science, Department of the Navy, Washington, D.C. Bassot, J.-M. (1966). On the comparative morphology of some luminous organs. I n “Bioluminescence in Progress” (Eds F. H. Johnson and Y . Haneda), pp. 557-610. Princeton University Press, New Jersey. Beamish, P. C. (19714. Acoustic scattering from zooplankton organisms. I n “Biological Sound Scattering in the Ocean” (Ed. G. B. Farquhar), pp. 474-475. Maury Center for Ocean Science, Department of the Navy, Washington, D.C. Beamish, P. C. (1971b). Quantitative measurements of acoustic scattering from zooplankton organisms. Deep-sea Rwearch, 18, 811-822. Beasley, T. M., Heyraud, M., Higgo, J . J. W., Cherry, R. D. and Fowler, S. W. (1978). zl’JPo and zl’JPbin zooplankton faecal pellets. Marine Biology, 44, 325-328.

Beaudouin, J. (1973). Zooplancton du Golfe de Gascogne (plateau continental) en 1971. Annabs biologiques, 28, (1971), 70-72. Beaudouin, J. (1974). Zooplancton du Golfe de Gascogne (plateau continental) en 1972. Annalw biologiques, 29 (1972), 61-63. Beaudouin, J. (1977). Euphausiaces et mysidaces du Golfe de Gascogne (plateau continental) en 1971. Annales biologiques, 32 (1975), 67-69. Bedard, J. (1969). Feeding of the least, crested, and parakeet auklets around St. Lawrence Island, Alaska. Canadian Journal of Zoology, 47, 1025-1050. Beklemishev, K. V. (1960). Southern atmospheric cyclones and the whale feeding grounds in the Antarctic. Nature, London, 187, 530-531. Belloni, S., Cattaneo, R. and Pessani, D. (1976a). Determinazione di acidi grassi in Meganyctiphanes norvegica (Sam, 1857) (Crustacea Euphausiacea) del Mar Ligure. Bollettino dei musei e degli instituti biologici della ( R . ) Universita d i Cfenova,44, 103-112. Belloni, S., Cattaneo, R., Orlando, P. and Pessani, D. (1976b). Alcune oonsiderazione sul contenuto in metalli pesanti in Meganyctiphanes normegica (Sara, 1857) (Crustacea Euphausiacea) nel Mar Ligure. Bolleltino dei musei e degli instituti biologici della ( R . ) Universita d i Benova, 44, 113-133.

REFERENOES

557

Benayoun, G., Fowler, S. W. and Oregioni, B. (1974). Flux of cadmium through euphausiids. Marine Biology, 27, 206-212. Bentz, K. L. M. (1976). Gill arch morphology of the Cape hakes Merlucciua capensis Cast. and M . paradoxua Franca. Fisheries Bulletin of South Africa, NO.8, 17-22. Berkes, F. (1975). Some aspects of feeding mechanisms of euphausiid crustaceans. Crustaceana, 29, 266-270. Berkes, F. (1976). Ecology of euphausiids in the Gulf of St. Lawrence. Journal of the Fisheries Research Board of Canada, 33, 1894-1905. Berkes, F. (19774. Production of the euphausiid crustacean ThysanoEasa raechii in the Gulf of St. Lawrence. Journal of the Fisheries Research Board of Canada, 34, 443-446. Berkes, F. (197713). Ecological significance of the neglecta form of ThysanoEasa inermis (Krayer) (Euphausiacea). Crustaceam, 33, 39-46. Binet, D. (1976). Contribution a 1’6cologie de quelques taxons du zooplancton de CBte d’Ivoire. IV, EuphausiacBs. Document acienti$que de Centre Recherche Ochnographique, Abidjan, 7 , 86-90. Bishop, D. G., James, D. G. and Olley, J. (1976). Lipid composition of slender tuna (Allothunnuafallai) as related to lipid composition of their feed (Nyctiphanes auatralis). Joumzal of the Fisheries Rmearch Board of Canada, 33, 1156-1161.

Blackburn, M. (1977). Studies on pelagic animal biomasses. I n “Oceanic Sound Scattering Prediction” (Eds N. R. Andersen and B. J. Zahuranec), Marine Science, Vol. 5, pp. 283-299. Plenum Press, New York and London. Blackburn, M., Laura, R. M., Owen, R. W. and Zeitzschel, B. (1970). Seasonal and areal changes in standing stocks of phytoplankton, zooplankton,and micronekton in the eastern tropical Pacific. Marine Biology, 7 , 14-31. Bliznichenko, T. E., Degtereva, A. A., Drobysheva, S. S., Nesterova, V. N., Rossova, E. Y. and Ryzhov, V. M. (1978). Results of plankton investigations in the Norwegian and Barents Seas in 1976. A n n a b biologiques, 33 (1976), 64-56.

Boden, B. P. (1960). The post naupliar stages of the crustacean Euphauaia pueifica. Transactione,of the Alnerican Microscopical Society, 69, 373-386. Boden, B. P. (1951). The egg and larval stages of Nyctiphanea simplex, a euphausiid crustacean from California. Proceedings of the Zoological Society of London, 121, 615-527.

Boden, B. P. (1964). The euphausiid crustaceans of southern African waters. Tranaactiona of the Royal Society of South Africa, 84, 181-243. Boden, B. P. (1955). Euphausiacea of the Benguela Current. “Di8covery” Reports, 27, 337-376.

Boden, B. P. and Brinton, E. (1957). The euphausiid crustaceans Thysanopoda aeqwtlis Hansen and Thysanopoda subaeqwzlia Boden, their taxonomy and distribution in the Pacific. Linznology and Oceamgraphy, 2, 337-341. Boden, B. P., Johnson, M. W. and Brinton, E. (1955). The Euphausiacea (Crustacea) of the North Pacific. Bulletin of the Scripps Imtitution of Oceanography. 0, 287-400.

Bogdanov, M. A. and Solyanik, Ye. V. (1970). Variations in the amount of ocean. Oceanology, MOSCOW, 11, 699-702. Bogdanov, M. A. and Solyankin, Ye. V. (1970). Variations in the amount of Euphausia superba Dana around South Georgia Island in relation to the hydrological regime. Oceanology, Moscow, 10, 542-547.

658

THE BIOLOQY OF EUPHAUSJIDS

Bogdanov, M. A., Oradovskiy, S. G., Solyanik, Ye. V. and Khvatskiy, N. V. (1969). On the frontal zone of the Scotia Sea. Oceanology, Moscow, 9, 777-783. Bottino, N. R. (1972). Fatty acid exchange among trophic levels of the Ross Sea; phytoplankton, copepods and euphausiids. Antarctic Journal, 7 , 178-179. Bottino, N. R. (1973). The lipids of Antarctic Krill. Presence of waxes in Euphau8thcrystdlorophiaa but not in Euphawia auperba 1ipids.FederationProceedings. Federation of American Societies for Experimental Biology, 32, 562 Abs. Bottino, N. R. (1974). The fatty acids of Antarctic phytoplankton and euphausiids. Fatty acid exchange among trophic levels of the Ross Sea. Marine Biology, 27, 197-204. Bottino, N. R. (1975). Lipid composition of two species of Antarctic krill: Euphauaia auperba and E . crystallorophiaa. Comparative Biochemiatry and Phy~iology,50B, 479-484. Boucher, J. and Thiriot, A. (1972). Zooplancton et micronecton estivaux des deuxcents premiers metres en MBditerranBe occidentale. Marine Biology, 15, 47-56.

Boyd, C. M. (1973). Small scale spatial patterns of marine zooplknkton examined by an electronic in Situ zooplankton detecting device. Netherlade Journal of Sea Research, 7 , 103-111. Boyd, S. H., Wiebe, P. H. and Cox, J. L. (1978). Limits of Nematoacelis megalops in the northwestern Atlantic in relation to Gulf Stream cold core rings. 11. Physiological and biochemical effects of expatriation. Journal of Marine Research, 36, 143-159. Boyden, C. R., Crothers, J. H., Little, C. and Mettam, C. (1977). The intertidal invertebrate fauna of the Severn Estuary. Field Studies, 4, 477-554. Bradbury, M. G., Abbott, D. P., Bovbjerg, R. V., Mariscal, R. N., Fielding, W. C., Barber, R. T., Pearse, V. B., Proctor, S. J., Ogden, J. C., Wourms, J. P., Taylor, Jr., L. R., Christofferson, J. G., Christofferson, J. P., McPhearson, R. M., Wynne, M. J. and Stromborg Jr., P. M. (1970). Studies on the fauna associated with the deep scattering layers in the equatorial Indian Ocean, conducted on R/V Te Vega during October and November, 1964. In “Biological Sound Scattering in the Ocean” (Ed. G. B. Farquhar), pp. 409-452. Maury Center for Ocean Science, Department of the Navy, Washington, D.C. Bradford, J. M. (1972). Systematicsand ecology of New Zealand central east coast plankton sampled at Kaikoura. New Zealad Oceanographic Imtitute Menzoir, No. 64, 87 pp. Brattelid, T. E. and Matthews, J. B. L. (1978). Studies on the deep-water pelagic community of Korsforden, Western Norway. The dry weight and caloric content of Euchaeta norvegica (Copepoda), Boremy& arctica (Mysidacea), and Meganyctiphanes norvegkm (Euphausiacea). Sarsia, 63, 203-21 1. Bright, T., Ferrari, F., Martin, D., and Franceschini, G. A. (1972). Effects of a total solar eclipse on the vertical distribution of certain oceanic zooplanktern. Limnology and Oceanography. 17, 2 9 6 3 0 1 . Brinton, E. (1962). The distribution of Pacific euphausiids. Bulktin of the Sm*pps Inatitution of Oceanography, 8, 51-270. Brinton, E. (1966). Remarks on Euphausiacean phylogeny. I n “Symposium on Crustacea”, Vol. 1, pp. 255-259. Marine Biological Association of India. Brinton, E. (19678). Distributional Atlas of Euphausiacea (Crustacea) in the California Current region. Part I. California Cooperative Oceanic FbherieS Investigatiom, Atlas 5 , 1-275.

REFERENCES

659

Brintoa, E. (1967b). Vertical migration and avoidance capability of euphausiids in %heCalifornia Current. Lirnnology and Oceanography, 12, 451-483. Brinton, E. (1973). Distributional atlas of Euphausiacea (Crustacea) in the California Current region, Part 11. California Cooperative Oceanic Fisheriea Innuestigationa, Atlas 18, 1-336. Brintan, E. (1975). Euphausiids of southeast Asian waters. Naga Report, 4 (a), 1-287. Brinton, E. (1976). Population biology of Euphausia paci,fica off southern California. Fishery Bulbtin, National Oceanic Atmospheric Administration of the United States, 74, 733-762. Brinton, E. (1978). Observations of spermatophores attached to pleopods of preserved male euphausiids. Crustaceans, 35, 241-248. Brinton, E. (1979). Parameters relating to the distributions of planktonic organisms, especially euphausiids in the eastern tropical Pacific. Progress in Oceanography, 8, 125-189. Brinton, E. and Gopalakrishnan, K. (1973). The distribution of Indian Ocean euphausiids. I n “Biology of the Indian Ocean” (Ed. B. Zeitzschel), pp. 357382. Chapman and Hall, London. Brinton, E. and Wyllie, J. G. (1976). Distributional atlas of euphausiid growth stages off southern California, 1953 through 1956. California Cooperative Oceanic Fkheries Investigationa, Atlas 24, 1-289. Brown, S. G. (1968). Feeding of sei whales at South Georgia. Norsk Hvalfangattide&, No. 6, 118-125. Bulycheva, N. P., Lozbin, L. I., Majstruk, P. N. and Marova, M. S. (1977). Microflora of the “Ocean” krill paste as a quality index. Cfigienai sanitariya, 3, 102-104. Burkholder, P. R., Mandelli, E. F. and Centeno, P. (1967). Some chemical properties of Munida gregaria and E u p h k superba. Journal of Agricultural and Food Chemistry, 15, 718-720. Bykov, V. P. (1976). Biological value of Antarctic krill and possibility of their utilization for human consumption. I n “Oceanology International 75, Into Deeper Waters”, pp. 361-363. BPS Exhibitions, London (U.K.). Casanova, B. (1970). RBpartition bathymBtrique des euphausiacBs dens le bassin occidental de la MBditerranBe. Revue des Travaux. Imtitut des Ptches mritime.9. 34, 205-219. Casanova, B. (1972). C16 de dBtermination des larves calyptopis des euphausiacBs de MBditerranBe. Cmcstaceana, 22, 178-180. Casanova, B. (1974). Les euphausiacBs de MBditerranBe (Systematique et dBveloppement larvaire. Biogeographie et biologie). Thitse pour Docteur Essciences Naturelles, L’Universite de Provence, Aix-Marseille, 380 pp. Casanova, B., Ducret, F. and Rampal, J. (1973). Zooplancton de MBditerranBe orientale et de Mer Rouge (Chaetognathes EuphausiacBs, PtBropodes). Rapport et procl-verbaux des rduniom. G m h & o n intermtionale pour a mar Mdditerranke, 21, 515-519. l’exploration sckntifiue de l Casanova, J.-P. (1977). La f a m e pelagique profonde (zooplancton et micronecton) de la province Atlanto-MBditerranBe. Aspects taxonomiques, biologiques et zoogeographiques. These pour Docteur Es-sciences Naturelles, L’Universite de Provence, Aix-Marseille, 455 pp. Casanova-Soulier, B. (1968). C1B de d6termination des larves furcilia des euphausiac6s de la MBditerranBe. Rapport et procb-verbaux des rduniom. Commission interncltionalepour l‘expbration scientijique de la mar Mdditerranb, 19,627-529.

560

THE BIOLOGY OF EUPHAUSIIDS

Casanova-Soulier, B. (19704. Les euphausiachs de MBditerran6e. Rapport et prochverbaux des rkunions. Commission. internationale pour l’exploration scientifque de la mer Mkditerranke, 20, p. 417. Casanova-Soulier, B. (1970b). Les rassemblements d‘euphausiac6s en MBditerran6e. Rapport et pro&-verbaux des rdunions. Commission internationale pour l’exploration scientijlque de la mer Mdditerranhe, 20, 435-437. Chekunova, V. I. and Rynkova, T. I. (1974).Energy requirements of the Antarctic krill, Euphausia superba Dane. Oceanology, Moscow, 14, 434-440. Chen, C.-S., Gau, S.-W. and Yan, T.-R. (1978). Studies on the proteases of Antarctic krill, Euphausia superba: activities and stability. Journal of the Chinese Biochemical Society, 7, 78-87. Cherry, R. D., Higgo, J. J. W. and Fowler, S. W. (1978). Zooplankton fecal pellets and element residence times in the ocean. Nature, London, 274, 246-248. Childress, J. J. (1975). The respiratory rates of midwater crustaceans as a function of depth of occurrence and relation to the oxygen minimum layer off southern California. Deep-sea Research, 50A, 787-799. Childress, J. J . (1977). Physiological approaches to the biology of midwater organisms. In “Oceanic Sound Scattering Prediction” (Eds N. R. Andersen and B. J. Zahuranec), Marine Science, Vol. 5, pp. 301-324. Plenum Press, New York and London. Childress, J. J. and Nygaard, M. (1974). Chemical composition and buoyancy of midwater crustaceans as function of depth of occurrence off southern California. Marine Biology, 27, 225-238. Clarke, A. (1976). Some observations on krill (Euphausia superba Dana) maintained alive in the laboratory. Bulletin of the British Antarctic Sarvey, No. 43, 111-118. Clarke, T. A. (1978). Die1 feeding patterns of 16 species of mesopelagic fishes from Hawaiian waters. Fishery Bulletin, National Oceanic Atmospheric Administration of the United Statw, 76, 495-513. Clutter, R. I. (1969). The microdistribution and social behaviour of some pelagic mysid shrimps. Journal of Experimental Marine Biology and Ecology, 3, 125-155. Costanzo, G. and Guglielmo, L. (1976a). Diagnostic value of the thelycum in euphausiids, I. Mediterranean species (first note). Crustaceana, 31, 45-52. Costanzo, G. and Guglielmo, L. (1976b). Diagnostic value of the thelycum in euphausiids, I. Mediterranean species (second note). Crustaceana, 31, 178-180. Costanzo, G. and Guglielmo, L. (1977). Sur l’importance du thelycum dans la systematique des esphces voisines du genre Thysanoessa (Euphausiacea). Rapports et procis-verbaux des rkunions. CommisGon internationale pour l’exploration scientijlque de la mer Mdditerranke, 24, 129-131. Cox, J. L. (1971). Uptake, assimilation, and loss of DDT residues by Euphausia paci$ca, a euphausiid shrimp. Pishery Bulletin, National Oceanic Atmospheric Administration of the United States, 69, 627-633. Cox, S. J. (1975). Shore stranding of Meganyctiphanes norvegica (M. Sars). Estuarine and Coastal Marine Science, 3, 483-484. Cram, D. L. and Schiilein, F. H. (1974). Observations on surface-shoaling Cape hake off South West Africa. Journal du Conseil, 35, 272-275. Croce, N.D. and Chiarabini, A. (1971). A suction pump for sampling mid-water and bottom organisms in the sea. Deep-sea Research, 18, 851-854.

REFERENOES

561

Cross, F. A., Fowler, S. W., Dean, J. M., Small, L. F. and Osterberg, C. L. (1968). Distribution of ? Z n in tissues of two ‘marine crustaceans determined by auto-radiography. Journal of the Fisheries Research Board of Canada, 25, 2461-2466. Culkin, F. and Morris, R . J. (1969). The fatty acids of some marine crustaceans. Deep-sea Research, 16, 109-116. Cushing, D. H. and Richardson, I. D. (1956). A record of plankton on the echosounder. Journal of the Marine Biological Association of the United Kingdom, 35, 231-240. Cutshall, N. H., Naidu, J. R. and Pearcy, W. G. (1977). Zinc and cadmium in the Pacific hake Merluccius productus off the western U.S. coast. Marine Biology, 44, 195-201. Czeczuga, B. and Klyszejko, B. (1977). Carotenoids in Euphausia superba (Dana, 1850) (Euphausiacea)-a representative of criles of the Antarctic waters. Bulletin de I’Academie polonake des sciences, SLrie des Sciences biologiques, 25, 597-600. Day, D. S. (1971). Macrozooplankton and small nekton in the coastal waters off Vancouver Island (Canada) and Washington, spring and fall of 1963. National Oceanic Atmospheric Administration of the United States, Special ScientiJic Report-Fisheries, No. 619, 1-94. De Bhaldraithe, P. (1973). A record of the ellobiopsid Thahsomyces fagei (Boschma, 1948) parasitic on the euphausiid Meganyctiphanes norvegica (M. Sars, 1856). Bulletin of the Zoological Museum of the University of Anwterdam, 3, 69-71. De Decker, A. (1973). Agulhas Bank plankton. I n “The Biology of the Indian Ocean” (Ed. B. Zeitzschel), pp.189-219. Chapman and Hall, London. Degtyaryova, A. A., Drobysheva, S. S., Nesterova, V. N., Rossova, E. Y., Ryzhov, V. M. and Soboleva, M. S. (1976). Plankton investigations in the Norwegian and Barents Seas in 1974. Annabs biologiques, 31 (1974), 54-56. Donaldson, H. A. (1975). Vertical distribution and feeding of sergestid shrimps (Decapoda : Natantia) collected near Bermuda. Marine Biology, 31, 37-50. Dragovich, A. (1969). Review of studies of tuna food in the Atlantic Ocean. United States Fish and Wildlife Service, Special ScientiJic Report-Fisheries, NO. 593, 1-21. Dragovich, A. (1971). The food of skipjack and yellowfin tunas in the Atlantic Ocean. Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 68, 445-460. Drobysheva, S. S. and Soboleva, M. S. (1976). The abundance of euphausiids from the Barents Sea in 1954-1972. Trudy Vsesoyuznogo Nauchno-issledovatel’ skogo Instituta Morskogo Rybnogo Khozyaistva i OkeanograJii (Trudy V N I R O ) , 110, 81-84. Drobysheva, S. S., Degtyaryova, A. A., Nesterova, V. N., Rossova, E. Y. and Ryzhov, V. M. (1977). Results of plankton investigations in the Norwegian and Barents Seas in 1975. Annulea biologiques, 32, (1975), 55-56. Ebeling, A. W., Cailliet, G. M., Ibara, R . M., DeWitt, F. A. and Brown, D. W. (1971). Pelagic communities and soundscattering off Santa Barbara, California. I n “Biological Sound Scattering in the Ocean” (Ed. G. B. Farquhar), pp. 1-19. Maury Center for Ocean Science, Department of the Navy, Washington, D.C. Eddie, G. C. (1977). The harvesting of krill. Food and Agricultural Organization of the United Nations, United Nations Development Programme, Rome, Report GLO/S0/77/2, 1-76.

562

THE BIOLOGY OF EUPHAUSIIDS

Egan, W. G. and Conrad, J. E. (1975). Summer abundance and ecology of zooplankton in the Gulf’ Stream. Biological Bulletin, Marine Biological Laboratory, Woods Hole, Mass., 149, 492-505. Einarsson, H. (1942). Notes on Euphausiacea 1-111. On the systematic value of the spermatheca, on sexual dimorphism in Nemtoscelis and on the male in Bentheuphausia. VidenskabeligeMeddeleleer fra Dansk naturhistorisk Forening i Kabenhavn, 106, 263-286. Einarsson, H. (1945). Euphausiacea I. North Atlantic species. Dana Report, NO.27, 1-185. Elder, D. L. and Fowler, S. W. (1977). Polychlorinated biphenyls: penetration into the deep ocean by zooplankton fecal pellet transport. Science, 197, 459-461. Elofsson, R. and Dahl, E. (1970). The optic neuropiles and chiasmata of Crustacea. Zeitschrift fiir Zellforschung, 107, 343-360. El-Sayed, S. Z. (1978). Primary productivity and estimates of potential yields of the Southern Ocean. I n “Polar research, to the present and the future” (Ed. M. A. McWhinnie), pp. 141-160. American Association for the Advancement of Science. El-Sayed, S. Z. and McWhinnie, M. A. (1979). Antarctic krill: protein of the last frontier. Oceanus, 22, 13-20. Endo, Y. and Komaki, Y. (1979). Larval stages of euphausiids with descriptions of those of Thysanogssa longipes Brandt. Bulletin of the Japan Sea Regional Fisheries Research Laboratory, No. 30, 97-1 10. Enomoto, Y. (1972). On the food chain in the Yellow Sea and in the East China Sea. Bulletin of the Tokai Regional Fisheries Reaearch Laboratory, 69, 65-73. Everson, I. (1976). Antarctic krill: a reappraisal of its distribution. Polar Record, 18, 15-23. Everson, I. (1977). The living resources of the southern ocean. Food and Agricultural Organization of the United Nations, United Nations Development Programme, Rcme, Report GLO/SO/77/1, 1-156. Fairbridge, R. W., ed. (1966). “The Encyclopedia of Oceanography”, 1021 pp. Reinhold Publishing Corporation, New York. Farquhar, G. B., ed. (1971). “Biological Sound Scattering in the Ocean”, 629 pp. Maury Center for Ocean Science, Department of the Navy, Washington, D.C. Farquhar, G. B. (1977). Biological sound scattering in the oceans: a review. I n “Oceanic Sound Scattering Prediction” (Eds N. R. Andersen and B. J. Zahuranec),Marine Science, Vol. 5, pp. 493-527. Plenum Press, New York and London. Fedotova, N. A. (1976). Composition and distribution of macroplankton in the Gulf of Tatary. Trudy Vsesoyuznogo Nauchno-~sledovate~’8kogo Instituta Morskogo Rybnogo Khozyaistva i Okeanografii (Trudy VNIRO), 110, 28-31. Fedotova, N. I., Baranov, V. S., Mikhailov, S. K. and Skurikhin, I. M. (1977). The amino acid composition of the “Ocean” krill paste as affected by heat treatment. Voprosy pitaniya, 3, 84-88. Ferguson, C. F. and Raymont, J. K. B. (1974). Biochemical studies on marine zooplankton. XII. Further investigations on Euphausia superba Dana. Journal of the Marine Biological Association of the United Kingdom, 54, 719-725. Ferrari, F. D. (1976). The significance of the response of pelagic marine animals to solar eclipses. Deep-sea Research, 23, 653-654.

REFERENCES

563

Field, L. H. (1969). The biology of Notophryxus ZaMralis (Isopoda: Epicaridea), parasitic on the euphausiid Nematoscelis dimilia. J o u r d of Parasitology, 55, 1271-1277. Fischer, S. (1974). Der Krill (Euphausia superba) und anders Nahrungs-resewen im Gebeit der Antarktis. Protokoll zur Fischerietechnik, 13, 226-288. Fischer, W. (1976): Tagesperiodische Wanderungen des Antarktischen Krill. Informations Fischwirtschaft, 23, 90-92. Fisher, L. R. (1967).Pigments of euphausiid eyes. I n “Symposium on Crustacea”, Vol. 3, pp. 1074-1080. Marine Biological Association of India. Flechtenmacher, W., Schreiber, W., Christians, 0. and Roschke, N. (1976). Die Verarbeitung von Krill. Informations Fiachwirtachaft, 23, 188-196. Fleminger, A., Isaacs, J. D. and Wyllie, J. G. (1974). Zooplankton biomass measurements from CalCOFI cruises of July, 1955 to 1959 and remarks on comparison with results from October, January and April cruises of 1955 to 1959. California Oceanic Fisheries Investigations, Atlas 12, 1-1 18. Folsom, T. R. and Beasley, T. M. (1963). Contributions from the alpha emitter, polonium-210, to the natural radiation environment of the marine organisms. In “Radioactive Contamination of the Marine Environment”, pp. 625-632. International Atomic Energy Agency, Vienna. Forsyth, D. C. T. and Jones, L. T. (1966). Swarming of Thysanoessa longicaudata (Kreyer) (Crustacea, Euphausiacea) in the Shetland Islands. Nature, London, 212, 1467-1468. Fowler, S. W. and Benayoun, G . (1976).Selenium kinetics in marine zooplankton. Marim Science Communications, New York, 2, 43-67. Fowler, S. W. and Heyraud, M. (1976). Flux of plutonium through marine biota. Rapport et procl-verbaux des rtfunions. Commission internationale pour l’exploratwn de lamer Me‘diterrantfe,23, 161-162. Fowler, S. W. and Small, L. F. (1972). Sinking rates of euphausiid fecal pellets. Lintnology and Oceanogrqhy, 17, 293-296. Fowler, S. W., Small, L. F. and Dean, J. M. (1969). Matabolism of zinc-65 in euphausiids. I n “Symposium on Radioecology” (Eds D. J. Nelson and F. C. Evans), CONF-670503,pp. 399-41 1. United States Department of Commerce, Springfield, Virginia. Fowler, S. W., Small, L. F. and Dean, J. M. (1970). Distribution of ingested zinc65 in the tissues of some marine crustaceans. Journal of the Fisheries Research Board of Canada, 27, 1051-1058. Fowler, S. W., Benayoun, G. and Small, L. F. (1971a). Experimental studies on feeding, growth and assimilation in a Mediterranean euphausiid. T h a h s i a Jugoslavica, 7, 35-47. Fowler, S. W., Small, L. F. and Dean, J. M. (1971b). Experimental studies on elimination of zinc-65, cesium- 137 and cerium-144 by euphausiids. Marine Biology, 8, 224-231. Fowler, S. W., Small, L. F. and Keckes, S. (1971~).Effects of temperature and size on moulting of euphausiid crustaceans. Marine Biology, 11, 48-5 1. Fowler, S. W., Small, L. F. and La Rosa, J. (1972). The role of euphausiid molts in the transport of radionuclides in the sea. Rapport et procl-verbaux dea rtfunions. Commission internationale pour l’exploration acientifiue de la mer Mt?diterran&, 21, 291-292. Fowler, S. W., Heyraud, M., Small, L. F. and Benayoun, G. (1973). Flux of 141Cethrough a euphausiid crustacean. Marine Biology, 21, 317-325.

564

THE BIOLOGY OF EUPHAUSIIDS

Foxton, P. and Roe, H. S. J. (1974). Observations on the nocturnal feeding of some mesopelagic decapod Crustacea. Marine Biology, 28, 37-49. Franqueville, C. (1970). Etude comparative de macroplancton en MBditerranhe nord-occidentale par pIong6es en soucoupe S P 360, et pbches au chalut pelagique. Marine Biology, 5, 172-179. Franqueville, C. (197 1). Macroplancton profond (Invert6brBs) de la MBditerranbe nord-occidentale. Tethys, 3, 11-56. Fraser, F. C. (1936). On the development and distribution of the young stages of krill (Euphausia superba). “Discoveq” Reports, 14, 3-192. Fraser, J. H. (1970). Scottish plankton collections 1969 : zooplankton. Annalee biologiques, 26 (1969), 112. Fraser, J. H. and Seaton, D. D. (1969). Scottish plankton collections, 1968. Annales biologiques, 25 (1968), 91-93. Freytag, G. (1977). Lottechnische Beobachtungen beim Fang von Krill und potentiellen Nutzfischen der Antarktis. ProtokoUe zur Fischereitechnik, 64, 54-68. Friedl, W. A. (1971a). Sonic scattering and probable causes in two areas of Pug& Sound. I n “Biological Sound Scattering in the Ocean” (Ed. G. B. Farquhar), pp. 527-548. Maury Center for Ocean Science, Department of the Navy, Washington, D.C. Friedl, W. A. (1971b). The relative sampling performance of 6- and 10-foot Isaacs-Kidd midwater trawls. Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 69, 427-432. Fujika, T., Yamamoto, T., Yamagi, I. and Shipematsu, T. (1969). Ash, iron, manganese content of marine plankton. Journal of the Japamse Chemical Society, 90, 680-686. Translation series of the Fisheries Research Board of Canada, No. 2557, 19?3, 1-25. Fukushima, S. (1976). Food intake of Pacific saury. Bulbtin of the Japanese Society of Scientific Fisheries, 42, 1189 Fulton, J. (1976). Frozen marine zooplankton as food for larval and juvenile fishes. Manuscript Report Series of the Fisheries Research Board of Canada, NO. 1390, 1-23. Galt, J. H. and Whisler, H. C. (1970). Differentiation of flagellated spores in ThalaPsomyces ellobiopsid parasite of marine Crustacea. Archiv f a r Mikrobiologk, 71, 29.5303. Gardner, G. A. (1977). Analysis of zooplankton population fluctuations in the Strait of Georgia, British Columbia. Journal of the Fisheries Research Board of Canada, 34, 1196-1206. Gaskin, D. E. (1976). The evolution, zoogeography and ecology of Cetace’a. Oceanography and Marine Biology, an Annual Review, 14, 247-346. Geiger, S. R., Rodriguez, K. and Murillo, M. M. (1968). Euphausiacea of the Arctic Ocean and its peripheral seas. Bulletin of the Southern California Academy of Sciences, 67, 69-79. George, R. Y. and Marum, J. P. (1974). The effects of hydrostatic pressure on living aquatic organisms. 111. Behaviour and tolerance of euplanktonic organisms to increased hydrostatic pressure. Internationale Revue der Qesamten Hydrobiologie u. Hydrographie, 59, 175-180. Gilfllan, E. S. (1972a). Seasonal and latitudinal effects of the responses of Euphausia pacifica Hansen (Crustacea) to experimental changes of temperature and salinity. I n “Biological Oceanography of the Northern North Pacific Ocean”, (Eds A. Y. Takenouti et al.), pp. 443-463. Idemitsu Shoten, Tokyo.

REFERENCES

565

Gillillan, E. S. (1972b). Reactions of Euphausiapacifica Hamen (Crustacea) from oceanic, mixed oceanic-coastal and coastal waters of British Columbia to experimental changes in temperature and salinity. Journal of Experirnental Marine Biology and Ecology, 10, 29-40. Gilfillan,E. S. (1976).Effects of water-body characteristics other than temperature and salinity on respiration in Eupha&pacijka. Marine Biology, 38, 305-313. Gjdsaeter, J. (1973). The food of the myctophid fish, Benthosema glaciale (Reinhardt) from western Norway. Sarsia, 52, 53-68. Glover, R. S. (1952). Continuous plankton records: the Euphausiacea of the north-eastern Atlantic and the North Sea, 1946-1948. Bulletin of Marine Ecology, 3 , 185-214. Gopalakrishnan, K. (1973). Developmental and growth studies of the euphausiid Nematoscelis di&ilis (Crustacea) based on rearing. Bulletin of the Scripps Institution of Oceanography, 20, 1-87. Gopalakrishnan, K. (1974). Zoogeography of the genus Newtoscelis (Crustacea, Euphausiacea). Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 72, 1039-1074. Gopalakrishnan, K. (1975). Biology and taxonomy of the genus Newmtoscelis (Crustacea, Euphausiacea). Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 7 3 , 797-814. Gopalakrishnan, K. and Brinton, E. (1969). Preliminary observations on the distribution of Euphausiacea from the International Indian Ocean Expedition. Bulletin of the National Institute of Sciences of India, No. 38, 694-611. Grantham, G. J. (1976). The Antarctic Krill. Manuscript published by Unilever Research, Aberdeen, Scotland, 86 pp. Grantham, G. J. (1977).The utilization of krill. Food and Agricultural Organization of the United Nations, United Nations Development P r o g r a m , Report GLO/SO/77/3, 1-61. Greenlaw, C. F. (1977). Backscattering spectra of preserved zooplankton. Journal of the Acoustical Society of America, 62, 44-52. Greenlaw, C. F. (1979). Acoustical estimation of zooplankton populations. Limnology and Oceanography, 24, 226-242. Greze, V. N., Gordejava, K. T. and Shmeleva, A. A. (1969). Distribution of zooplankton and biological structure in the tropical Atlantic. I n “Proceedings of the Symposium on the Oceanography and Fisheries Resources of the Tropical Atlantic”, pp. 85-90. UNESCO, France. Groisman, M. Ya., Karpenko, E. A. and Stepanov, G. N. (1969). The experimental fishing of krill in the Scotia Sea. Trudy Y ~ ~ ~ u z Namhmwm g o Issledovatel’skogo Instituta Morskogo Rybnogo Khozyaktva i Okeanografii (Trudy VNIRO), 66, 276-283. National Lending Library for Science and Technology, Boston Spa, Yorkshire, RTS 5592. Gros, P. and Cochard, J.-C. (1978). Biologie de Nyctiphanes couchii (Crustacea, de Euphausiacea) dans le secteur nord du Golfe de Gascogne. Ann& 1’Institut Ocdanographique,54, 25-46. Gudkov, V. M. (1962). Relationship between the distribution of zooplankton, sea birds and baleen whales. Trudy Instituta okeanobgii, Akademiya nauk, SSSR,58, 298-313, Translation of the United States Naval Oceanographic Office, N O 0 T-7 (1974). Guglielmo, L. (1969). Spiaggia#mentidi eufausiacei lung0 la costa Messinese dello Strettodal Dicembre 1968 a1 Dicembre 1969. Bolletino d i pesca, piscicoltura e idrobiologia, 24, 7 1-77.

566

THE BIOLOQY OF EUPEAUSIIDS

Gulielmo, L. and Costanzo, G. (1977). Diagnostic value of the thelycum in euphausiids, 11. Oceanic species. Genus Thysanopodu Milne-Edwards, 1830. Crustaceana, 33, 275-283. Gulland, J. A. (1970). The development of the resources of the Antarctic seas. I n “Antarctic Ecology” (Ed M. W. Holdgate), Vol. 1, pp. 217-223. Academic Press, London and New York. Haedrich, R. L. and Henderson, N. R. (1974). Pelagic €ood of Coryphaenoidee amnatua, a deep benthic rattail. Deep-sea Research, 21, 739-744. Halcrow, W., Mackay, D. W. and Thornton, I. (1973). The distribution of trace metals and fauna in the Firth of Clyde in relation to the disposal of sewage sludge. Journal of the Marine Biological Association of the United Kingdom, 53, 721-739. Hansen, H. J. (1905). Further notes on the Schizopoda. Bulktin du Musde Ocdanographiquede Monaco, No. 42, 1-32, Hansen, H. J. (1910). The Schizopoda of the Siboga Expedition. Reports of the Siboga Expedition, 37, 1-123. Hansen, R. P. (1969). The occurrence of phytanic acid in Antarctic krill. (Euphauaia superba). Australian Journal of Science, 32, 160-161. Hansen, R. P. and Meiklen, S. M. (1970). Isoprenoid fatty acids in Antarctic krill (Euphaueia superba). Journal of the Science of Food and Agriculture, 21, 203-206. Harbison, G. R., Madin, L. P. and Swanberg, N. R. (1978). On the natural history and distribution of oceanic ctenophores. Deep-sea Research, 25, 233-256. Harding, G. C. H. (1974). The food of deep-sea copepods. Journal of the Marine Biological Association of the United Kingdom, 54, 141-155. Herding, G. C. H. (1977). Surface area of the euphausiid Thysano&sa rmchii and its relation to body length, weight, and respiration. Journal of the Fisheries Research Board of Canada, 34, 225-231. Hart, T. J. (1942). Phytoplankton periodicity in Antarctic surface waters. ‘‘DiBcovery” Reports, 21, 26 1-356. Harvey, B. J. (1977). Circulation and dioptric apparatus in the photophores of Euphauaia pacifiilica: some ultra-structural observations. Canadian Journal of Zoology, 55, 884-889. Haury, L. R. (1973). Sampling bias of a Longhurst-Hardy plankton recorder. Limnology and Oceanography, 18, 500-506. Haury, L. R. (1976a). Small-scale pattern of a California Current zooplankton assemblage. Marine Biology, 37, 137-157. Haury, L. R. (1976b). A comparison of zooplankton patterns in the California Current and North Pacific central gyre. Marine Biology, 37, 159-167. Haury, L. R., Wiebe, P. H. and Boyd, S. H. (1976). Longhurst-Hardy plankton record-: their design and use to minimize bias. Deep-sea Research, 23, 1217-1229. Heegaard, P. (1948). Larval stages of Meganyctiphanes (Euphausiacea) and some general phylogenetic remarks. M&blser fra Konzmissionen for Danmrks Fiskeri og Havunders0gelser,Series Plankton, 5 ( l ) , 1-27. Hempel, I. (1979). Vertical distribution of eggs and nauplii of krill (Euphauaia superba) south of Elephant Island. Meeresforschung, Reporta on Marine Research, 27, 119-123. Hempel, I. and Hempel, G. (1978). Larval krill (Eupham*a auperba) in the plankton and neuston samples of the German Antarctic Expedition 1975/76. Meeresforschung, Repor& on Marine Raearch, 26, 206-2 16.

REFERENCES

567

Henderson, J. T. (1927). A note on the effect of temperature on the cardiac rhythm of certain schizopods. Journal of Experimental Biology, 5 , 135-137. Herring, P. J. (1972). Depth distribution of the carotenoid pigments and lipids of some oceanic animals. 1. Mixed zooplankton, copepods and euphausiids. Journal of the Marine Biological Association of the United Kingdom, 52, 179-189. Herring, P. J. and Locket, N. A. (1978). The luminescence and photophores of euphausiid crustaceans. Journal of Zoology, 186, 431-462. Hesthagen, I. H. (1973). Diurnal and seasonal variations in the near-bottom fauna-the hyperbenthos-in one of the deeper channels of the Kieler Bucht (Western Baltic). Kieler Meeresforschungen, 29, 116-140. Hesthagen, I. H. and Gjermundsen, B. (1978). The replicability of sampling the hyperbenthic regions by means of Beyer’s 50cm epibenthic closing net. Meeresforschung, Reports on Marine Research, 26, 1-10. Heyraud, M. and Cherry, R. D. (1979). Polonium-210 and lead-210 in marine food chains. Marine Biology, 52, 227-236. Heyraud, M., Fowler, S . , Beasley, T. and Cherry, R. (1976a). Distribution du 2lOPo dans le zooplancton marin. Rapport et procl-verbaux des rduniona. Commission internationale pour l’exploration sckntijlque de la rner Miditenande, 23, 123-124. Heyraud, M., Fowler, S. W., Beasley, T. M. and Cherry, R. D. (197613). Polonium210 in euphausiids: a detailed study. Marine Biology, 34, 127-136. Higgo, J. J. W., Cherry, R. D. Heyraud, M. and Fowler, S. W. (1977). Rapid removal of plutonium from the oceanic surface layer by zooplankton faecal pellets. Nature, London, 266, 623-625. Hirano, T. (1975). On the distribution and seasonal variation of homarine in some marine invertebrates. Bulletin of the Japanese Society of Scientific Fisheries, 41, 1047-1 05 1. Hobson, E. S. and Chess, J. R. (1976). Trophic interactions among fishes and zooplankters near shore a t Santa Catalina Island, California. Fkhery Bulletin, National Oceanic Atmospheric Administration of the United States, 74, 567-598. Holden, M. J. and Tucker, R. N. (1974). The food of Raja clavata Linnaeus 1758, Raja montagui Fowler 1910, Raja naevus Muller and Henle 1841 and Raja brachyura Lafont 1873 in British waters. Journal du Conseil, 35, 189-193. Holdgate, M. W., ed. (1970). “Antarctic Ecology”, Vol. 1, 604 pp. Academic Press, London and New York. Hollingshead, K. W. and Corey, S. (1974). Aspects of the life history of Meganyctiphanes norvegica (M. Sars), Crustacea (Euphausiacea) in Passamaquoddy Bay. Canadian Journal of Zoology, 52, 495-505. Holtzman, R. B. (1969). Concentrations of naturally occurring radionuclides, 226Ra, ZlOPb, and 2loPo in aquatic fauna. I n “Symposium on Radioecology” (Eds D. J. Nelson and F. C. Evans), CONF-670503, pp. 535-546. United States Department of Commerce, Springfield, Virginia. Hong, S. Y. (1969). The euphausiid crustaceans of Korean waters. I n “The Kuroshio (a Symposium on the Japan Current)” (Ed. J. C. Marr), pp. 291300. University of Hawaii Press. Hopkins, C. C. E. and Gulliksen, B. (1978). Diurnal vertical migration and zooplankton-epibenthos relationships in a north Norwegian fjord. I n “Physiology and Behaviour of Marine Organisms” (Eds D. S. McLusky and A. J. Berry), pp. 271-280. Pergamon Press, Oxford.

568

THE BIOLOGY OF EUPHAUSIIDS

Hopkins, C. C. E., Falk-Petersen, S., Tande, K. and Eilertsen, H. C. (1978). A preliminary study of zooplankton sound scattering layers in Balsfjorden : structure, energetics and migrations. Sarsia, 63, 255-264. Hopkins, T. L. (1968). Carbon and nitrogen content of fresh and preserved Nemtoscelia di&ilis, a euphausiid crustacean. Journal du Conseil, 31, 300-304.

Hopkins, T. L. and Baird, R. C. (1973). Diet of the hatchetfish Sternoptyz diapbna. Marine Biology, 21, 34-46. Hopkins, T. L. and Baird, R. C. (1977). Aspects of the feeding ecology of oceanic midwater fishes. I n “Oceanic Sound Scattering Prediction” (Eds N. R. Andersen and B. J. Zahuranec), Marine Science, Vol. 5, pp. 325-360. Plenum Press, New York and London. Hopkins, T. L., Baird, R. C. and Milliken, D. M. (1973). A messenger-operated closing trawl. Limnology and Oceanography, 18, 488-490. Hu, V. J. H. (1978). Relationships between vertical migation and diet in four species of euphausiids. Limnology and Oceanography, 23, 296-306. Idler, D. R. and Wiseman, P. (1971). Sterols of Crustacea. International Journal of Biochemistry, 2, 91-98. Ikeda, T. (1970). Relationship between respiration rate and body size in marine plankton animals as a function of the temperature of habitat. Bulletin of the Faculty of Fisheries of Hokkaido University, 21, 91-112. Ikeda, T. (1972). Chemical composition and nutrition of zooplankton in the Bering Sea. I n “Biological oceanography of the Northern North Pacific Ocean” (Eds A. Y. Takenouti et al.),pp. 433-442. Idemitsu Shoten, Tokyo. Ikeda, T. (1977a). The effect of laboratory conditions on the extrapolation of experimental measurements to the ecology of marine zooplankton. 11. Effect of oxygen saturation on the respiration rate. Bulletin of the Plankton Society of Japan, 24, 19-28. Ikeda, T. (1977b). The effect of laboratory conditions on the extrapolation of experimental measurements to the ecology of marine zooplankton. IV. Changes in respiration and excretion rates of boreal zooplankton species maintained under fed and starved conditions. Marine Biology, 41, 241-252. Isaacs, J. D., Fleminger, A. and Miller, J. K. (1969). Distributional atlas of zooplankton biomass in the California Current region: spring and fall 19551959. California Cooperative Oceanic Fisheries Investigations, Atlas 10, 1-252.

Isaacs, J. D., Fleminger, A. and Miller, J. K. (1971). Distributional atlas of zooplankton biomass in the California Current region : winter 1955-1959. California Cooperative Oceanic Fisheries Investigations, Atlas 14, 1-122. Itoh, K. (1970). A consideration on feeding habits of planktonic copepods in relation to the structure of their oral parts. Bulletin of the Plankton Society of Japan, 17, 1-10. Ivanov, B. G. (1969). On the luminescence of the Antarctic krill ( E u p h u k a superba). Okeanologija, Moscow, 9, 505-506. Ivanov, B. G. (1970). On the biology of the Antarctic krill Euphausia superba. Marine Biology, 7, 340-351. Jagersten, G.(1972). “Evolution of the Metazoan Life Cycle”, 282 pp. Academic Press, London and New York. Jahn, F., Lukovicz, M. and Loerz, R. (1978). Erfahrungen mit Krillmehl im Forellenfutter. Praktischer Fuetterungsversuch. Fischerei Teichwirt, 29, 67-68.

REFERENCES

669

James, B. M. (1970). Euphausiacean Crustacea. Oceanographic Studies of Texas A . & M . University, 1, 205-229. James, P. T. (1973). Distribution of dimorphic males of three species of Nematoscelis (Euphausiacea). Marine Biology, 19. 341-347. James, P. T. (1977). The comparative morphology of the thelycum in the Euphauaia gibba species group. Deep-sea Research, 24, 1037-1047. Jawed, M. (1969). Body nitrogen and nitrogenous excretion in Neornysis rayii Murdoch and Euphauaia paci$ca Hansen. Lirnnology and Oceanography, 14, 748-754.

Jawed, M. (1973). Ammonia excretion by zooplankton and its significance to primary productivity during summer. Marine Biology, 23, 115-120. Jennings, C. D. (1977). SsFe in Pacific Ocean plankton. Marine Biology, 44, 223-226.

Jennings, J. B. and Halverson, L. G. (1971). Variations in subcuticular acid phosphatase activity during the molting cycle of the euphausiid crustacean, Thyaanoessarmchii (M. Sam) Hansen. Journal of Marine Research, 29,13%139. Jerde, C. W. and Lasker, R. (1966). Molting of euphausiid shrimps: shipboard observations. Lirnnology and Oceanography, 11, 120-124. Jillett, J. B. (1971). Zooplankton and hydrology of Hauraki Gulf, New Zealand. Memoir of the New Zealand Oceanographic Inatitute, No. 53, 1-103. Jillett, J. B. (1976). Zooplankton associations off Otago Peninsula, south-eastern Journal of New Zealand, related to different water masses. New Z&nd Marine and Freshwater Reeearch, 10, 543-557. Jillett, J. B. and Mitchell, S. F. (1973). Hydrological and biological observations in Dusky Sound, south-western New Zealand. I n “Oceanography of the South Pacific” (Ed. R. Fraser), pp. 419-427. New Zealand National Commission for UNESCO, Wellington. Johannessen, Per J. (1977). Ecological studies on the deep-water pelagic community of Korsfjorden, Western Norway. Studies of sampling with a Longhurst-Hardy Plankton Recorder. Sarah, 63, 85-91. John, D. D. (1936). The southern species of the genus Euphctusia. “ficovery” Reports, 14, 193-324. Jones, D. A. (1971). A new light trap for plankton. I n “Fourth European Marine Biological Symposium” (Ed. D. J. Crisp), pp. 487-493. Cambridge University Press, Cambridge. Jones, L. T. (1969). Continuous plankton records: studies on the zooplankton east of Newfoundland and Labrador, with particular reference to the euphausiid Thysano&sa longicaudata (Kreyer). Bulletin of Marine Ecology, 6 , 275-300. Jorgensen, G. and Matthews, J. B. L. (1975). Ecological studies on the deepwater pelagic community of Korsfjorden, Western Norway. Population dynamics of six species of euphausiids in 1968 and 1969. Sarsia, 59, 67-84. Kagei, N., h n o , K. and Kihata, M. (1978). On the examination against the parasites of Antarctic krill, Euphauaia swperba. Scienti$c Reports of the Whales Research Institute of Tokyo, No. 30, 311-313. Kanaeva, I. P. and Pavlov, V. Ya. (1976). To the biology of Nyctiphanes simplex Hansen. Trudy V8esoyuznogo Nauchno-iasledovatel’skogoInstituta Morskogo Rybnogo Khozyaistva i Okeanogra$i (Trudy V N I R O ) , 110, 90-96. Karedin, E. P. (196%). Preliminary report on the composition and distribution of macroplankton (euphausiids and others) in the Kuroshio Area. Summer 1965-Winter 1966. Izveetiya Tikhookeanskogo Nauchno-~sledovatel’8kogo Instituta Rybnogo Khozyaktva i Okeunograjii, 68, 125-141.

570

THE BIOLOQY OF ECJPHAUSIIDS

Karedin, E. P. (196913). The seasonal variations of macroplankton in the Kuroshio Area. Izwestiya Tikhookeanskogo Nauchno-issledowatel‘skogoInstituta Rybnogo Khozyaistwa i Okeanografii, 68, 142-166. Karedin, E. P. (1971). About identity between Nematoscelis megalops G. 0. Sara, 1885, N. dificilis Hansen, 1911 (Euphausiacea, Crustacea) and validity of distinguishing N . diBcilis Hansen. Izwestiya Tikhookeanskogo Nauchnoissledowatel’skogoInstituta Rybnogo Khozyaistwa i Okeanogra$i, 75, 121-129. Kashkin, N. I. (1976). On macroplankton from the southwest part of the Barents Sea. Trudy V8esoyuznogo Nauchno-issledowatel’skogo Instituta Morskogo Rybnogo Khozyaiatwa i Okeanografii (Trudy V N I R O ) , 110, 40-47. Kawakami, T., Hamabe, M. and Saito, R. (1973). A preliminary note on the ecology of the ommastrephid squid Nototodaw sloani sloani (Gray) in New Zealand waters. Bulletin of the Tokai Regional Fisheries Research Laboratory, 76, 53-69. Kawamura, A. (1970). Food of sei whale taken by Japanese whaling expedition in the Antarctic season 1967-68. Scientifi Reports of the Whales Research Institute of Tokyo, No. 22, 127-152. Kawamura, A. (1973). Food and feeding of sei whale caught in the waters south of 40°N in the North Pacific. Scientific Reports of the Whales Research Institute of Tokyo, No. 25, 219-236. Kawamura, A. (1974a). Food and feeding ecology in the southern sei whale. Scientific Reporh of the Whales Research Institute of Tokyo, No. 26, 25-144. Kawamura, A. (1974b). Euphausiids from the viewpoint of food cycle in the seas. Bulletin of the Japanese Society of Fisheries Oceanography, 24, 39-49. Kawamura, A. (1977). On the food of Bryde’s whales caught in the South Pacific and Indian Oceans. Scientific Reports of the W h a h Research Institute of Tokyo, No. 29, 49-58. Kayama, M.,Ikeda, Y. and Komaki, Y. (1976). Studies on the lipids of marine zooplankton, with special reference t o wax ester distribution in crustaceans and its in viwo formation. Yu-Kagaku, 25, 329-334. KefkeS, S., Fowler, S. W. and Small, L. F. (1972). Flux of different forms of loaRuthrough a marine zooplankter. Marine Biology, 13, 94-99. Kelle, W. (1978). Zur frage von Beschaedigungsraten des Krills in der Kommerziellen Fischerei. Informations Fischurirtschaft, 25, 59-61. Kelly, J. A. and Dragovich, A. (1968). Occurrence of macroplankton in Tampa Bay, Florida and the adjacent Gulf of Mexico. Fishery Bulletin of the United States Fisheries and Wildlife Service, No. 66, 209-221. Kelley, M. D., Lukaschewsky, S. and Anderson, C. G. (1978). Bacterial flora of Antarctic krill (Euphamia superba) and some of their enzymatic properties. Journal of Food Science, 43, 1196-1197. Kennedy, V. S. (1976). Arsenic concentrations in some co-existing marine organisms from Newfoundland and Labrador. Journal of the Fisheries Research Board of Canada, 33, 1388-1393. Khan, M.A. and Williamson, D. I. (1970). Seasonal changes in the distribution of Chaetognatha and other plankton in the eastern Irish Sea. Journal of Experimental Marine Biology and Ecology, 5, 285-303. Kikuchi, T. and Kano, S. (1975). Analysis of the metal elements in Euphau-~ia superba by UHF plasma spectra analyzer. Journal of the Tokyo University of Fisheries, 62, 11-17.

REFERENCES

571

Kils, U. (1979).Performance of Antarctic krill Euphawria auperba, at different levels of oxygen saturation. Results of the 2 German Antarctic Expedition 1977178.Meeresforschung. Reports on Marine Reaearch, 27, 35-48. King, F. D.and Packard, T. T. (1975a).The effect of hydrostatic pressure on respiratory electron transport system activity in marine zooplankton. Deep-sea Reaearch, 22, 99-105. King, F. D. and Packard, T. T. (1975b).Respiration and the activity of the respiratory electron transport system in marine zooplankton. Limnology an& ‘Oceanography,20, 849-854. Kinzer, J. (1967). Die Verbreitung des Zooplanktons in Echostreuschichten extrem sauerstoffarmen Wassers. Umrrchau, 22, 733-734. Kinzer, J. (1969a). On the quantitative distribution of zooplankton in deep scattering layers. Deep-sea Reaearch, 16, 117-125. Kinzer, J. (1969b).Quantitative distribution of zooplankton in surface waters of the Gulf of Guinea during August and September, 1963. I n “Proceedings of the Symposium on Oceanography and Fisheries Resources of the Tropical Atlantic”, pp. 231-240. UNESCO/FAO, Paris. Kinzer, J. (1971). On the contribution of euphausiids and other plankton organisma to deep scattering layers in the eastern north Atlantic. I n “Biological Sound Scattering in the Ocean” (Ed. G. B. Farquhar), pp. 476-489. Maury Center for Ocean Science, Department of the Navy, Washington, D.C. h e r , J. (1977).Observations on feeding habits of the mesopelagic fish Benthoaema g h i a l e (Myctophidae) off N.W. Africa. I n “Oceanic Sound Scattering Prediction” (Eds N. R. Andersen and B. J. Zahuranec), Marine Science, Vol. 5, pp. 381-392. Plenum Press, New York and London. Kiortsis, V. (1974). Quelques consid6rations sur 1’6cologie du zooplancton en Mer EgBe. RBpartition g6ographiques et variations saisonnihres. Rapport et procb-verbaux dea rduniona. Cornmiasion internationale pour l’exploration acientifiue de la m r Mdditerrande, 22, 139-141. Klyashtorin, L. B. and Yarzhombek, A. A. (1973).On energy losses for active movement of planktonic organisms. Okeanologija, Moecow, 13, 697-703. Knauer, G.A.and Martin, J. H.(1972).Mercury in a marine pelagic food chain. Limnology and Oceanography, 17, 868-876. Knight, M. D. (1973).The nauplius 11, metanauplius and calyptopis stages of Thyaanopoda tricuepidata Milne-Edwards (Euphausiacea). Fishery Bulletin, National Oceanic Atmospheric Adminiatration of the United States, 71, 53-67. Knight, M. D. (1975). The larval development of Pacific Euphauaia gibboidea (Euphausiacea). Fishery Bulletin, National Oceanic Atm8pheric Administration of the United States, 73, 145-168. Knight, M. D. (1976). The larval development of Euphaush aanzoi Torelli (Crustacea, Euphausiacea). Bulletin of Marine Science, 26, 538-557. Knight, M. D. (1978).Larval development of Euphausia falhx Hansen (Crustacean: Euphausiacea) with a comparison of larval morphology within the E . gibboidea species group. Bulletin of Marine Science, 28,255-281. Knox, G. A. (1970).Biological oceanography of the South Pacific. I n “Scientific Exploration of the South Pacific” (Ed. W. S. Wooster),pp. 155-182. National Academy of Sciences, Washington, D.C. Koch, H. and Lambert, K. (1976). Investigations by the German Democratic Republic on blue ling (Molva byrkehnge Walb.) in September, 1973 east of the Faroes. Annales biobgique8, 31 (1974),115-116.

572

THE BIOLOGY OF EUPHAUSIIDS

Kock, K.-H. and Neudecker, A. (1977). Krillfaenge im Atlantischen Sektor der Antarktis. Informations Fischdrtschaft, 24, 8-12. Kock, K.-H. and Stein, M. (1978). Krill and hydrographic conditions off the Antarctic Peninsula. Meeresforschung, Reports on Marine Research, 26, 79-95. Koehler, R. (1911). Isopodes nouveaux de la famille des Dajid6s provenant des Campagnes de la “Princesse Alice”. Bulletin de Z’lnstitut Ochnographique, Monaco, 196, 1-34. Komaki, Y. (1967). On the surface swarming of euphausiid crustaceans. Pacific Science, 21, 433-448. Komaki, Y. (1970). On the parasitic organisms in a krill, Euphausia sirnilis, from Suruga Bay. Journal of the Oceanographical Society of Japan, 26, 283-295. Koops, H., Tiews, K., Beck, H. and Gropp, J . (1977). Ersatz von Fischmel durch Alkanhefe und Krillmehl im Forellenfutter. Informations Fischwirtschaft, 24. 21-22. Korabel’nikov, L. V. (1975). Some zoogeographical characteristics of the Antarctic Balleny Islands. Vestnik Zoologiskk 3, 35-41. Koyama, T., Nakamura, M. and Okamoto, H. (1974). An experiment with a large model of surface-midwater trawl for krill. Bulletin of the Tokai Regional Fisheries Research Laboratory, No. 78, 1-10. Kravitz, M. J., Pearcy, W. G. and Guin, M. P. (1976). Food of five species of cooccurring flatfishes on Oregon’s continental shelf. Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 74, 984-990. Kryuchkova, M. I. and Makarov, 0. E. (1969). Technochemical characteristics of krill. Trudy Vsesoyuznogo Nauchno-issledovatel’skogo Instituta Morskogo Rybnogo Khozyaistva i Okeanografii (Trudy V N I R O ) , 66, 295-298. National Lending Library for Science and Technology, Boston Spa, Yorkshire, RTS 5594. Kuenzler, E. J. (1969a). Elimination of iodine, cobalt, iron and zinc by marine zooplankton. I n “Symposium on Radioecology” (Eds D. J. Nelson and F. C. Evans), CONF-670503, pp. 462-473. United States Department of Commerce, Springfield, Virginia. Kuenzler, E. J. (1969b). Elimination and transport of cobalt by marine zooplankton. I n “Symposium on Radioecology” (Eds D. J. Nelson and F. C. Evans), CONF-670503, pp. 483-492. United States Department of Commerce, Springfield, Virginia. Kulka, D. W. and Corey, S. (1978). The life history of Thysano&sa inermis (Krcayer)in the Bay of Fundy. Canadian Journ,al of Zoology, 56, 492-506. Kuwano, K. and Mitamura, T. (1977). On the Antarctic krill protein concentrate (KPC). I. Manufacture of the KPC by isopropyl alcohol method and its chemical composition. Bulletin of the Japanese Society of Scientific Fisheries, 43, 559-565. Lacroix, G. and Bourget, E. (1973). The forms inermis and neglecta of Thysanoessa inermis (Krcayer, 1849) (Euphausiacea) in the southwestern part of the Gulf of St. Lawrence, E. Canada. Crustaceana, 24, 298-302. LagardBre, J.-P. (1972). Recherches sur l’alimentation des crevettes de la pente continentale Marocaine. Tethys, 3, 655-675. LagardBre, J.-P. (1973). Donn6es sur la biologie et sur l’alimentation de Dichelopalzdalus bonnieri (Crustac6-Natantia) dam le Golfe de Gascogne. Tethys, 5, 165-166.

REFERENOES

573

Lagardbre, J.-P. (19774. Recherches sur le regime alimentaire et le comportement predateur des decapodes benthiques de la pente continentale de 1’Atlantique nord oriental Golfe de Gascogne et Maroc). I n “Biology of Benthic Organisms” (Eds B. F. Keegan, P. 0. Ceidigh and P. J. S. Boaden), pp. 397408. Pergamon Press, Oxford. LagardBre, J.-P. (1977b). Recherches sur la distribution verticale et, sur l’alimentation des crustaces decapodes benthiques de la pente continentale du Golfe de Oascogne. Analyse des groupements carcinologiques. Bulletin du Centre d’Etudes et de Recherches scientifiques Biamitz, 11,367-440. La Rosa, J. (1976). A simple system for recovering zooplanktonic faecal pellets in quantity. Deep-sea Research, 23, 996-997. Lasker, R. (1964). Moulting frequency of a deep sea crustacean, Euphuusia pacifica. Nature, London, 203, 96. Lasker, R. (1966). Feeding, growth, respiration, and carbon utilization of a euphausiid shrimp. Journal of the Fkheries Research Board of Canada, 23, 1291-1317.

Lasker, R. and Theilacker, G. H. (1965). Maintenance of euphausiid shrimps in the laboratory. firnnology and Oceanography, 10, 287-288. Latogurakii, V. I. (1972). Euphausia superba Dana as a food item of Notothenia rossi mrmorata. Trudy Atlantic nauchno-Issfivatel’skogoInatituta Morskogo Rybnogo Khozyaistva i Okeanografii (Trudy Atlantic NIRO), 42, 167-169. Latogurskii, V. I., Naumov, A. G. and Pervushin, A. S. (1975). Antarctic divergence, the biology and distribution of krill. Trudy Atlantic nauchno-issledovatel’skogo Inatituta Morskogo Rybnogo Khozyaistva i Okeanografii (Trudy Atlantic NIRO), 58, 77-88. Leatherland, T. M., Burton, J. D., Culkin, F., McCartney, M. J. and Morris, R. J. (1973). Concentrations of some trace metals in pelagic organisms and of mercury in northeast Atlantic Ocean water. Deep-sea Research, 20, 679-686. Lee, R. F. (1974). Lipids of zooplankton from Bute Inlet, British Columbia. Journal of the Fisheries Research Board of Canada, 31,1577-1682. Lee, R. F. and Hirota, J. (1973). Wax esters in tropical zooplankton and nekton and the geographical distribution of wax esters in marine copepods. Grnnology and Oceanography, 18, 227-239. Le Gall, J.-Y. and L’Herroux, M. (1972). Crustacbs pelagiques susceptibles de p6ches experimentales en Atlantique nord et en MBditeranee occidentale. Rapport scientifiue et technical de la centre national pour l’exploitation des O C ~ W , NO.8, 1-34. Legand, M., Bourret, P., Fourmanoir, P., Grandperrin, R., Gueredrat, J. A., Michel, A., Rancurel, P., Repelin, R. and Roger, C. (1972). Relations trophiques et distributions verticales en milieu pelagique dans l’Oc6an Pacifique intertropical. Cahiers 0.R.S.T.0.M.-Ochnographie, 10, 303-393. Legand, M., Bourret, P., Gueredrat, J. A., Michel, A., Repelin, R. and Roger, C. (1976). Aspects Bcologiques du plancton et du micronecton dans l’est de l’oc6an Indien. Journal of the Marine Biological Association of India, 16 (1974), 1-23.

Leinemann, M. and Christians, 0. (1977). Methods zur Pruefung der Gewebesaftlaessigkeit in Kochkrillfarce (KKF). I n f o m t i o n s Pischwirtschaft, 24, 124126.

Le Roux, A. (1973a). Observations sur le developpement larvaire de Nyctiphanes couchii (Crustacea: Euphausiacea) au laboratoire. Marine Biology, 22, 169166.

574

THE BIOLOGY OF EUPHAUSIIDS

Le Roux, A. (1973b). Influence de l’ablation des p6doncles oculaires sur Nyctiphanes couchii (Bell) (Crustac6, euphausiac6). C m p t e rendu hebdmadaire d a skances de l’academie des mienma, Pa&, 276, 1317-1320. Le Roux, A. (1974). Observations sur le d6veloppement larvaire de Meganyctip b m s norwegica (Crustacea: Euphausiacea) au laboratoire. Marine Biology, 26, 45-56.

Le Roux, A. (1976). Observations sur les larves de Nyctipbnes couchii et de Meganyctiphanes norvegica (Crustacea: Euphausiacea) recolt6es dans le Golfe du Morbihan. Cahiers de Biologie marine, 17,375-386. Leung, Y.-M. (1970). Key to the speciespf adult euphausiids of the Arctic Basin and its peripheral seas. I n “Taxonomic Guides to Arctic Zooplankton” (Eds Y.-M. Leung and H. A. Kobayashi), Vol. 1, pp. 39-46. Department of Biological Sciences, University of Southern California, Los hgeles. Lewis, J. B. (1955). Some larval euphausiids of the genus Stylocheiron from the Florida Current. Bulletin of Marine Science, Gulf and Caribbean, 5 , 190-202. Lewis, J. B. and Fish, A. G. (1969). Seasonal variation of the .zooplankton fauna of surface waters entering the Caribbean Sea at Barbados. Caribbean Journal of Science, 9, 1-24. Lie, U. (1907). The natural history of Hardangerfjord 8. Quantity and composition of the zooplankton, September, 1955-September, 1956. SarsicC, 30, 49-74.

Lindley, J. A. (1977). Continuous plankton records: the distribution of the Euphausiacea (Crustacea: Malacostraca) in the north Atlantic and the North Sea, 1966-1967. Journal of Biogeography, 4, 121-133. Lindley, J. A. (1978a). Populabion dynamics and production of euphausiids. I. Thysanoessa longicaudata in the North Atlantic Ocean. Marine Biology, 46, 121-130.

Lindley, J. A. (1978b). Continuous plankton records: the occurrence of apostome ciliates (Protozoa) on Euphausiacea in the North Atlantic Ocean and North Sea. Marine Biology, 46, 131-136. Lomakina, N. B. (1966). The euphausiid fauna of the Antarctic and notal regions. I n “Biological Reports of the Soviet Antarctic Expedition 1955-58”, Vol. 2, pp. 260-342. Israel Programme for Scientific Translation, Jerusalem. Lomakina, N. B. (1968). On the species independence of Thysanopoda 8Ubae4Wli8 Boden (Euphausiacea, Crustacea). Zoologicheskid Z h u d , Moscow, 47, 293295.

Lomakina, N. B. (1972). Phylogenetic relationships of the E u p h a ~ ~species h (Euphausiacea) of the South Ocean. Zoologischeskii Zhurnal, Moacow, 51, 812-820.

Lomakina, N. B. (1978). Euphausiids of the world oceans (Euphausiacea). I n “Determinations of the Fauna of the SSSR”, Vol. 118, pp. 1-222. Academy Nauk, SSSR. ~~~~~~~~, G., Naumov, A. G. and Lagunov, L. L. (1973). Prospects of the utilization of krill and other nonconventional resources of the world ocean. Journal of the Fisheries Research Board of Canada, 30, 2196-2201. Macdonald, A. G., Gilchrist, I. and Teal, J. M. (1972). Some observations on the tolerance of oceanic plankton to high hydrostatic pressure. Journal of the Marine Biological As8ociation of the United Kingdom, 52, 213-223. McEachran, J. D., Boesch, D. F. and Musick. J. A. (1976). Food division within two sympatric species-pairs of skates (Pisces: Rejidae). Marine Biology, 35, 301-317.

REFERENCES

575

McGowan, J. A. (1974). The nature of oceanic ecosystems. I n “The Biology of the Oceanic Pacific”, pp. 9-28. Oregon State University Press. McGowan, J. A. (1977). What regulates pelagic community structure in the Pacific? I n “Oceanic Sound Scattering Prediction” (Eds N. R. Andersen and B. J. Zahuranec), Marine Science, Vol. 5, pp. 423-443. Plenum Press, New York and London. Mackintosh, N. A. (1966). The swarming of krill and problems of estimating the standing stock. Norsk Hvalfangsttidende, No. 11, 113-1 16. Mackintosh, N. A. (1967).Maintenance of living Euphclusia superba and frequency of moults. Norsk Hvaljangsttidende, No. 5, 97-102. Mackintosh, N. A. (1970). Whales and krill in the twentieth century. I n “Antarctic Ecology” (Ed. M. W. Holdgate), Vol. 1, pp. 195-212. Academic Press, London and New York. Mackintosh, N. A. (1972). Life cycle of Antarctic Krill in relation t o ice and water conditions. “Discovey” Reporta, 36, 1-94. Mackintosh, N. A. (1973). Distribution of post-larval krill in the Antarctic. “DiBcovery” Reports, 36, 95-156. Mackintosh, N. A. (1974). Sizes of krill eaten by whales in the Antarctic. “Discovery” Reporta, 36, 157-178. McLaren, I. A. (1963). Effects of temperature upon growth of zooplankton, and the adaptive value of vertical migration. Journal of the Fisheries Research Board of Canada, 20, 685-727. McLaughlin, P. A. (1965). A redescription of the euphausiid crustacean, Nematoscelis d i f i i l i s Hansen, 1911. Crustaceana, 9, 41-44. McLellan, T. (1977). Feeding strategies of the macrourids. Deep-sea Research, 24, 1019-1036. Macpherson, E. (1978a). RBgimen alimentario de Micromsistius poutmaou (Risso, 1810) y Gadiculus argenteus argenteus Guichenot, 1850 (Pisces, Gadidae) en el Mediterrheo occidental. Investigacion Pesquera, 42, 305316. Macpherson, E. (1978b). RBgirnen alimentario de Phycis blennoides (Briinich) y Antonogadus mgalokynodon (Kolombatovic) (Pisces : Gadidae) en el Mediterrhea occidental. Investigacion Pesquera, 42, 455-466. Macquart-Moulin, C. and Leveau, M. (1968). RBcolte de macroplancton 6 partir de la bou6e laboratoire-juin 1966. Rapport et procl-verbaux des rdunions. Commission internationale pour E’exploration scientifZque de la w r MBditerran&, 19, 495-497. McWhinnie, M. A., Denys, C. and Schenborn, D. (1976). Biology of krill (Euphausia superba) and other Antarctic invertebrates. Antarctic Journal, United States, 11, 55-58. McWilliam, P. S. (1977). Further studies on plankton ecosystems in the eastern Indian Ocean. VI. Ecology of the Euphausiacea. Australian Journal of Marine and Freshwater Research, 28, 627-644. Makarov, R. R. (1970). Separate existence of various age groups of Antarctic krill. Information Bulletin of the Soviet Antarctic Expedition, 77, 547550. Makarov, R. R. (1971a). Some data on the composition of the population and peculiarities of reproduction of Euphausia supirba Dana. Oceanology, Moscow, 11, 73-80. Makarov, R. R. (1971b). Some pecularities of life cycles of euphausiids. Zoologicheakii Zhurnal, Moscow, 50,193-198.

576

THE BIOLOUY OF EUPHAUSIIDS

Makarov, R. R. (1972). The life cycle and features of distribution of Euphawia superba Dana. Trudy Vsesoyuznogo Nauchno-bsledovatel’skogoInstituta Morskogo Rybnogo Khozyaistva i 0kealzograflc.i (Trudy V N I R O ) , 77, 85-92. Makarov, R. R. (1973). Pecularities of the autumn state of Euphaauaia superba Dana population. Okeanologija, Moscow, 13, 334-338. Makarov, R. R. (1974a). Dominance of larval forms in eupiiousiid (Crustacea: Eucarida) ontogenesis. Marine Biology, 27, 93-100. Makarov, R. R. (1974b). Larvae of Euphauaia superba Dana in plankton from the Sea of Scotia. Trudy Vseaoyuznogo Nauchno-iasledovatel’skogo Imtituta Morskogo Rybnogo Khozyaiatva i OkeanograJii (Trudy) V N I R O , 99, 84-102. Makarov, R. R. (1975a). Vertical distribution of eggs and larvae in euphausiids of the north-eastern coast of the island of South Georgia. Okeanologia, Moscow, 15, 1101-1106. Makarov, R. R. (1975b). A study of repeated maturation of females (Eucarida, Euphausiacea). Zoologicheskii Zhumal, Moscow, 54,670-681. Makarov, R. R. (1976). To the biology ofreproduction of Euphausicc superba Dana (Crustacea: Euphausiacea). Trudy Vsesoyuznogo Nauchno-ksledovatel’skogo Imtituta Morskogo Rybnogo Khozyaistva i OkeanograJii (Trudy V N I RO), 110, 85-89. Makarov, R. R. (1977). Larval distribution and some aspects of reproduction ecology of Euphausia frigida Hansen, 1911 (Crustacea: Euphausiacea) in the southern part of the Scotia Sea. Okeanologija, Moscow, 17, 324-332. Makarov, R. R. and Maslennikov, V. V. (1975). Distribution and current drift in larvae of Euphawia superba in the Scotia Sea and adjacent waters. Soviet Journal of Marine Biology, 1, 37-43. Makarov, R. R. and Shevtsov, V. V. (1969). Antarctic krill. Prir&, No. 10, 81-83. Makarov, R. R. and Shevtsov, V. V. (1971). Some problems in the distribution and biology of Antarctic krill. Izdatel’stvo Nauka, Moscow, 81-87. National Marine Fisheries Service, Wmhington, D.C. Translation, No. 72-50077b (1972), 1-9. Makarov, R. R., Naumov, A. G. and Shevtsov, V. V. (1970). The biology and the distribution of the Antarctic krill. I n “Antarctic Ecology” (Ed. M. W. Holdgate), Vol. 1, pp. 173-176. Academic Press, London and New York. Markina, N. P. (1976). Quantitative distribution of macroplankton in the great Australian Bight with seasons. Trudy Vsesoyuznogo Nauchno-issledovatel’skogo Inatituta Morskogo Rybnogo Khozyaistva i OkeanograJii (Trudy V N I R O ) , 110, 21-27. Marlowe, C. J. and Miller, C. B. (1975). Patterns of vertical distribution and migration of zooplankton a t Ocean Station “P”. Limnology and Oceanography, 20, 824-844. Marr, J. W. S. (1962). The natural history and geography of the Antarctic krill (Euphausia mperba Dana).. “Dbcovery” Reports, 32, 33-464. Marshall, N. B. (1971). Swimbladder development and the life of deep-sea fishes. I n “Biological Sound Scattering in the Ocean” (Ed. G. B. Farquhar), pp. 69-73. Maury Center for Ocean Science, Department of the Navy, Washington, D.C. Mason, J. C. (1976). Response of underyearling coho salmon to supplemental feeding in a natural stream. Journal of Wildlife Management, 40, 775788.

REFERENCES

577

Mathew, K. J. (1971). Studies on the larval stages of Euphausiacea from the Indian Seas. I. Diagnostic characters of post-naupliar stages of Euphausia diomdeae Ortmann and E . distinguenda Hansen. Journal of the Marine Biological Association of India, 13, 52-60. Mathew, K. J. (1972). Studies on the larval stages of Euphausiacea from Indian Seas. 2. Description of post-naupliar stages of Stylocheiron carinaturn G. 0. Sars. Journal of the Marine. Biological Association of India, 14, 796-804. Mathew, K. J. (1975). Studies on the larval history of two species of Euphausiacea from the Indian Seas. Journal of the Marine Biological Association of India, 17, 1-16. Matthews, F. D., Damkaer, D. M., Knapp, L. W. and Collette, B. B. (1977). Food of western North Atlantic tunas (Thunnwr) and lancet fishes ( A k p i saurus). National Oceanic Atmspheric Administration of the United States, Technical Report NMFS SSRF-706, 1-19. Mathhews, J. B. L. (1973). Ecological studies on the deep-water pelagic community of Korsfjorden, western Norway. Population dynamics of Meganyctiphanes norvegica (Crustacea, Euphausiacea) in 1968 and 1969. Saraia, 54, 75-90. Matthews, J. B. L. and Bakke, J. L. W. (1977). Ecological studies on a deep-water pelagic community of Korsfjorden (western Norway). The search for 8 trophic pattern. HelgolLinder wissenschaftliche Meeresuntersuchungen, 30, 47-61. Matthews, J. B. L. and Hestad, L. (1977). Ecological studies on the deep-water pelagic community of Korsfjorden, western Norway. Length/weight relationships for some macroplanktonic organisms. Saraia, 63, 57-63. Mauchline, J. (1959). The development of the Euphausiacea (Crustacea) especially that of Megan,yctiphanes norvegica (M. Sara). Proceedings of the Zoological Society of London, 132, 627-639. Mauchline, J. ( 1960). The biology of the euphausiid crustacean, Meganyctiphanes norvegica (M. Sara). Proceedings of the Royal Society of Edinburgh, B . (Biology), 67, 141-179. Mauchline, J. (1965). The larval development of thB euphausiid Thysanoessa rmchii (M. Sars). Crustaceana, 99, 31-40. Mauchline, J. (1966). The biology of Thysanozssa rmchii (M. Sara), with a comparison of its diet with that of Meganyctiphanes norvegica (M. Sara). I n “Contemporary Studies in Marine Science” (Ed. H. Barnes), pp. 493-510. George Allen and Unwin, London. Mauchline, J. (1 967s). The feeding appendages of the Euphausiacea (Crustacea). Journal of Zoology, 153, 1-43. Mauchline, J. (196713). Volume and weight characteristics of Euphausiacea. Cwtaceana, 13, 241-248. Mauchline, J. (1971a). Crustacea, Euphausiacea: adults. Conseil International pour 1’Explorationde la Mer. Zooplankton Sheet No. 134, 1-8. Mauchline, J. (197lb). Crustacea, Euphausiacea: larvae. Comeil International pour I’Exploration de la Mer, Zooplankton Sheet Nos 135-137, 1-16. Mauchline, J. (1972). The biology of bathypelagic organisms, especially Crustacea. Deep-sea Research, 19, 753-780. Mauchline, J. ( 1 973). Assessing similarity between samples of plankton. Journal of the Marine Biological Association of India, 14 (1972), 26-41. Mauchline, J. (1976). The Hiatt growth diagram for Crustacea. Marine Biology, 35, 79-84.

678

THE BIOLOUY OF EUPHAUSIIDS

Mauchline, J. (19774. Growth of shrimps, crabs and lobsters-an assessment. Journal duComeil,37,162-169’. Mauchline, J. (197713). Estimating production of mid-water organisms. I n “Oceanic Sound Scattering Prediction” (Eds N. R. Andersen and B. J. Zahuranec), Marine Science, Vol. 5, pp. 361-400. Plenum Press, New York and London. Mauchline, J. (1977~).Growth and moulting of Crustacea, especially euphausiids. I n “Oceanic Sound Scattering Prediction” (Eds N. R. Andersen and B. J. Zahuranec), Marine Science, Vol. 5, pp. 401-422. Plenum Press, New York and London. Mauchline, J. (1977d). The integumental sensilla and glands of pelagic Crustacea. Journal of the Marine Biological A88OCiatiOn of the United Kingdom, 57, 973994. Mauchline, J. and Fisher, L. R. (1969). The biology of euphausiids. Advaneea i n Marine Biology, 7 , 1-454. Mauchline, J. and Nemoto, T. (1977). Integumental sensilla of diagnostic value in euphausiids. Journal of the Oceunographical Society of Japan, 33, 283-289. Mayer, R. (1967). Some data on the euphausiids of the Tirenian Sea and remarks on the diagnosis of Euphauaia krohnii Br. Travaux du Mueeum d’histoire naturelle W r . Antipa”, 7, 15-21. Maynard, S. D., Riggs, F. V. and Walters, J. F. (1975). Mesopelagic micronekton in Hawaiian waters: faunal composition, standing stock and die1 vertical migration. Fbhery Bulktin, National Oceanic Atmospheric Administration of the United s&h?8, 73, 726-735. Mayzaud, P. (19734. Respiration and nitrogen excretion of zooplankton 11. Studies of the metabolic characteristics of starved animals. Marine Biology, 21, 19-28. Mayzaud, P. (197313).Respiration et excr6tion azot6e du zooplankton. 111.Etude de l’influence des variations thermiques. Annales de l’Ina(itut ocdanogrmphique, Park, 49, 113-122. Mehl, J. A. P. (1969). Food of barracouta (Teleosti: Gempylidae) in eastern Cook Strait. New Zealand Journal of Marine and Freshwater Research, 3, 389-394. Meira, C. (1970). Contribuicao para o estudo dos eufausiaceos do arquipelago de Cab0 Verde. Notar, do Centro de Biologia Aqwtica Tropical, &boa, No. 19, 1-26. Meira, C. (1971). Nova esp6cie de Euphausiaceae parasitada por T h a ~ s o m y c e s fagei Boschma (Protozoa, Ellobiopsidae). Notas do centro de Biologia Aqwtica Tropical, Liaboa, No. 24, 1-9. Menzies, R. J. and Selvakumaran, M. (19748). The effects of hydrostatic pressure on living organisms. V. Eurybiotic environmental capacity as a factor in high pressure tolerance. Internationale Revue der gesamten Hydrobiologie u. Hydrographie, 59, 199-205. Menzies, R. J. and Selvakumaran, M. (197410): The effects of hydrostatic pressure on living aquatic organisms. VI.Organic complexity and hydrostatic pressure sensitivity. Internutionale Revue der geaamten Hydrobwlogie u. Hydrographie, 59, 207-212. Merrett, N. R. and Roe, H. S. J. (1974). Patterns and selectivity in the feeding of certain mesopelagic fishes. Marine Biology, 28, 115-126. Mezykowski, T. and Rakusa-Suszczewski, S. (1979). The circadian rhythms in Eupha& superba Dana and its carbohydrate metabolism. Meeresforschung, Reports on Marine Research, 27, 124-129.

REFERENOES

579

Michel, A. and Grandperrin, R. (1970). SBlection du chalut phlagique Isaacs-Kidd 10 pieds. Marine Biology, 6, 200-212. Micklukhina, A. P. (1969). Distribution and specific composition of plankton in the zone of the Pacific Drift. Izvestiya Tikhookeanskogo Nauchno-issledovatel’ skogo Inetituta Rybnogo Khozyaistva i Okeanografii, 68, 116-124. Mikhailovskii, Yu. A. (1969). Observations on krill in a zone of artificial illumination. Trudy Vsesoyuznogo Nauchno-issledovatel’akogo Instituta Morslcogo Rybnogo Khozyaktva i Okeanograjii (Trudy VNIRO), 66, 243-245. National Lending Library for Science and Technology, Boston Spa, Yorkshire, Translation RTS 6590. Miller, C. B. (1970). Some environmental consequences of vertical migration in marine zooplankton. Limnology and Oceanography, 15, 727-741. Miyake, Y., Sugimura, Y. and Mayeda, M. (1970). The uranium content and the activity ratio ag4U/238U in marine organisms and sea water in the western North Pacific. Journal of the OceanographicalSociety of Japan, 26, 123-129. Moguilevsky, A. and Gooday, A. J. (1977). Some observations on the vertical distribution and stomach contents of Grigantocypris muelleri Skogsberg 1920 (Ostracoda, Myodocopina). Sixth International Ostracod Symposium, 263270.

Mohr, H. (1976). Tageszeitlich bedingte Rhythmik im Verhalten von halbwiichsigem Krill ( E u p h w i a superba). Informtionem fiir die Fhchwirtschaft (Bundes-forschungsanstalt f fir Fwcherie, Hamburg), 23,132-134. Mohr, H. (1978). Beobachtungen iiber das Verhalten von antarktischen Krill in der Fangseison 1977178. Informations Fischwirtschaft, 25, 27-29. Mohr, H.and Fischer, W. (1977). Verhaltensbeobachtungenan Krillschwarmen. Protokolk zur Fkcherietechnik, 64, 39-53. Moiseev, P. A. (1970). Some aspects of the commercial use of the krill resources of the Antarctic Seas. I n “Antarctic Ecology” (Ed. M. W. Holdgate), Vol. 1, pp. 213-216. Academic Press, London and New York. Moore, E. and Sander, F. (1976). Quantitative and qualitative aspects of the zooplankton and breeding patterns of copepods at two Caribbean coral reef stations. Estuarine and Coastal Marine Science, 4, 689-607. Moore, E. and Sander, F. (1977). A study of the offshore zooplankton of the tropical western Atlantic near Barbados. Ophelia, 16, 77-96. Moroz, I. F. (1973). Hydrochemical conditions in areas inhabited by the Antarctic krill. Studies in Fisheries Biology and Oceanography, Vladivostock, 4, 36-43.

Morris, R. J. (1971). Variations in the fatty acid composition of oceanic euphausiids. Deep-sea Research, 18, 525-529. Morris, R. J. (1972a). The occurrence of wax esters in crustaceans from the northeast Atlantic Ocean. Marine Biology, 16, 102-107. Morris, R. J. (1972b). The preservation of some oceanic animals for lipid analysis. Jourml of the Fisheries Research Board of Canada, 29, 1303-1307. Morris, R. J. (1973). Relationships between the sex and degree of maturity of marine crustaceans and their lipid compositions. Journal of the Marine Biological Association of the United Kingdom, 53, 27-37. Morris, R. J. and Sargent, J. R. (1973). Studies on the lipid metabolism of eome oceanic crustaceans. Marine Biology, 22, 77-83. Mullin, M. M. (1969). Production of zooplankton in the ocean: the present status and problems. Oceanography and Marine Biology, an Annual Review, 7 , 293-314.

580

THE BIOLOGY OF EUPHAVSIIDS

Murano, M., Marumo, R., Nemoto, T. and Aizawa, Y. (1976). Vertical distribution of biomass of plankton and micronekton in the Kuroshio water off central Japan. Bulktin of the Plankton Society of Japan,, 23, 1-12. Myklebust, R. (1977). The membrane systems of the cardiac muscle cell of Meganyctiphanes norvegica (M. Sars), Euphausiacea, Crustacea. Cell and Tissue Research, 181, 339-347. Nagabhushanam, A. K. (1965). On the biology of the commoner gadoids in Manx waters. Journal of the Marine Biological Association of the United Kingdom, 45, 615-657.

Nagakura, K., Arima, S., Kurihara, M., Koga, T. and Fujita, T. (1974). Mercury content of whales. Bulletin of the Tokai Regional Fisheries Research Laboratory NO. 78, 41-46. Nakamura, E. (1967). Abundance and distribution of zooplankton in Hawaiian waters, 1955-56. United States Department of the Interior, Fish and Wildlife Service, Special Scientific Report-Fisheries, No. 544, 1-37. Nest, F. (1979). The vertical distribution of larval and adult krill (Euphamia auperba Dana) on a time station south of Elephant Island, South Shetlands. Meeresforschung, Reports on Marine Research, 27, 103-1 18. Naumov, A. G. and Permitin, Yu. E. (1973). On the trophic relations between Euphausia superba Dana and fish in the Southern Ocean with special reference to the Scotia Sea. T d y Vsesoyuznogo Nauchno-issledovatel’skogo Inatituta Morskogo Rybnogo Khozyaistva i Okeanografii (Trudy VNIRO), 93, 216-229. Naumov, A. G. and Ponomareva, L. A. (1967). Vertical distribution and diurnal migrations of the main representatives of zooplankton in the northern part of the Indian Ocean. T d y Instituta okeanologii Akademiya nauk SSSR, 64 (1964), 250-256. United States Naval Oceanographic Office Translation No. 329. Nemoto, T. (1959). Food of baleen whales with reference to whale movements. Scienti,jic Reports of the Whales Research Institute of Tokyo, No. 14, 149-290. Nemoto, T. (1966). Thysano&sa euphausiids, comparative morphology, d o morphosis and ecology. Scientific Reports of the Whales Research Institute of Tokyo, NO. 20, 109-155. Nemoto, T. (1969). EuphausiacBs’dans les couches intermt5diaires et profondes. Umi, 1, 50-55. Nemoto, T. (1970a). Feeding pattern of baleen whales in the ocean. I n “Marine Food Chains” (Ed. J. H. Steele), pp. 241-252. Oliver and Boyd, Edinburgh. Nemoto, T. (1970b). Chlorophyll pigments in digestive guts of zooplankton. Bulletin of the Plankton Society of Japan, 17, 50-55. Nemoto, T. (1972a). History of research into the food and feeding of euphausiids. Proceedings of the Royal Society of Edinburgh, 3. (Biology),73, 259-265. Nemoto, T. (1972b). Chlorophyll pigments in the stomach and gut of some macrozooplankton species. I n “Biological Oceanography of the Northern North Pacific Ocean” (Eds A. Y. Takenouti et al.), pp. 411-418. Idemitsu Shoten, Tokyo. Nemoto, T. (1977). Food and feeding structures of deep sea Thysanopoda euphausiids. I n “Oceanic Sound Scattering Prediction” (Eds N. R. Andersen and B. J. Zahuranec), Marine Science, Vol. 5, pp. 457-480. Plenum Press, New York and London. Nemoto, T. and Nasu, K. (1975). Present status of exploitation and biology of krill in the Antarctic. I n “Oceanology International 75, Into Deeper Waters”, pp. 353-360. BPS Exhibitions, London (U.K.).

REFERENOES

581

Nemoto, T. and Saijo, Y. (1968). Trace of chlorophyll pigments in stomachs of deep sea zooplankton.Journal of the OceanographicalSocietyof Japan, 24,4648. Nemoto, T., Ishikawa, K. and Kamada, K. (1969). A short note on the importance of the krill as a marine biological resource in relation to stock of baleen whales. Bulletin of the Japanese Society of Fisheries and Oceanography,Special Number in Convmewwration of Prof. Uda, 217-220. Nemoto, T., Kamada, K. and Ham, K. (1972). Fecundity of a euphausiid crustacean Nenzatoscelis di$cilis in the North Pacific Ocean. Marine Biology, 14, 41-47. Nemoto, T., Araki, M. and Brinton, E. (1973). Clinal variation in the frequency of one- and two-spinedforms of Thysanohsa inerrnis (Kreyer, 1849) (Euphausiacea) in the North Pacific. Cru&meana, 24, 318-322. Nemoto, T., Brinton, E. and Kamada, K. (1977). Reproduction and growth of deep sea Thysanopoda euphausiids. Bulletin of the Plankton Society of Japan, 24, 36-43.

Nepgen, C. S. de V. (1977). The biology of the hottentot Pachyrnetopon blochii (Val.) and the silverfish Argyrozona argyrozona (Val.) along the Cape southwest coast. Department of Industries, Republic of South Africa, Fisheries Branch Inveetiqdional Report, No. 106, 1-35. Nicholls, G. D., Curl, H. and Bowen, V. T. (1959). Spectrographic analyses of marine plankton. Limnology and Oceanography, 4, 472-478. Nikolaeva, G. G. and Ponomareva, L. A. (1973). Oxygen consumption by tropical euphausiids. Okeanologija, Moscow, 13, 493-496. Nival, P., Charm, R. and Palazzoli, I. (1973). Variation spatiale du t a m de respiration du zooplankton en MBditemn6e occidentale en Mars 1970: mission Mediprod I1 du Jean-Charcot. Annales de Z’Imtitut Ochnographique, Park, 49, 123-133. O’Connell, C. P. (1971). Variability of near-surface zooplankton off southern California, as shown by towed-pump sampling. Fishery Bulletin, National Oceanic Atrrwspheric Administration of the United States, 69, 681-697. Odate, K. (1977). On the feeding habits of the Pacific saury, Gololabis saira (Brevoort). Bulletin of the Tohoku Regional Fisheries Research Laboratory, 38, 76-88. Ogawa, K. (1977). The role of baoterial floc as food for zooplankton in the sea. BuUetin Japanese SoOiety Yor Scienti& Fisheries, 43, 395407. Ogi, H.and Tsujita, T. (1973). Preliminary examination of stomach contents of murres ( U k spp.) from the eastern Bering Sea and Bristol Bay, JuneAugust, 1970 and 1971. Japanese Journal of Ecology,2I, 201-209. Ohsumi, S.,Masaki, Y. and Kawamura, A. (1970). Stock of the Antarctic Minke whale. Scientifi Reporte of the Whales Reeearch I t i t U t e , Tokyo, No. 22, 75125.

Okada, S. and Taniguchi, A. (1971). Size rehtiomhip between salmon juveniles in shore waters and their prey animals. BuUetin of the Faculty of Fishrim, Hokkaido University, 2%, 30-36. Omori, M. (1969). Weight and chemical composition of some important oceanic zooplankton in the north Pacific Ocean. Marine Biology, 3, 4-10. Omori, M. (1978). Zooplankton fisheries of the world: a review. Marine Biology. 48, 199-205.

O’Riordan, C. E. (1969). A catalogue of the collection of Irish marine Crustacea in the National Museum of Ireland, 98 pp. Published by the Stationery Office, Dublin.

582

TIIE BIOLOGY OF EUPHAUSIIDS

Oritsland, T. (1970). Biology and population dynamics of Antarctic seals. I n “Antarctic Ecology”, (Ed. M.’ W. Holdgate), Vol. 1, pp. 361-366. Academic Press, London and New York. Oshima, T. (1969). A study of the first host of Aniaakia. Sawhin-Igaku, 24, 401404. Fisheries Research Board of Canada, Translation No. 1325, 1-12. Owens, T. G. and King, F. D. (1975). The measurement of respiratory electrontransport-system activity in marine zooplankton. Marine Biology, 30, 27-36. Owre, H. B. and Foyo, M. (1972). Studies on Caribbean zooplankton. Description of the program and results of the first cruise. Bulbtin of Marine Science, 22, 483-621.

Ozawa, K., Yamada, T., Kim, M. and Shimizu, T. (1968). Observations of patches of Euphausia superba. Journal of the Tokyo University of Fisheries, 9, 101-109.

Paranjape, M. A. (1967). Molting and respiration of euphausiids. Journal of the Fisheries Research Board of Canada, 24, 1229-1240. Parsons, T. R. and Lebrasseur, R. J. (1970). The availability of food to different trophic levels in the marine food chain. I n “Marine Food Chains” (Ed. J.H. Steele), pp. 325-343. Oliver and Boyd, Edinburgh. Parsons, T. R., Lebrasseur, R. J. and Fulton, J. D. (1967). Some observations on the dependence of zooplankton pazing on the cell size and concentration of phytoplankton blooms. Journal of the OceanographicalSociety of Japan, 23, 1C-17.

Pavillon, J.-F. (1978). Contribution to studies on the effect of ectocrine substances on the development of marine invertebrates. Absorption of labelled (14C or 3H) dissolved organic substances by the eggs and larvae of Euphausia krohnii. I n “Physiology and Behaviour of Marine Organisms” (Eds D. S. McLusky and A. J. Berry), pp. 149-154. Pergamon Press, London. Pavlov, V. Ye. (1969). The feeding of krill and some features of its behaviour. Trudy Vsesoyuznogo Nauchno-iasledovatel’skogoInstituta Mor8kogo Rybnogo Khozyaistva i Okeanografii (Trudy VNIRO), 86, 42-54. Pavlov, V. Ya. (1971a). On the physiology of feeding in Euphausia superba. Doklady Akademii nauk SSSR (Biological Sciences-Zoology), 196, 147-150. Pavlov, V. Ye. (1971b). On the qualitative composition of food for Euphausia superba Dana. Proceedings of the All-Union Reaearch Institute of Marine Fisher& and Oceanography, 86, 42-54. Fisheries Research Board of Canada, Translation Series, No. 2953, 1-16. Pavlov, V. Ya. (1974). On the relationship between the feeding habits and the behavioural characteristics of Euphausia superba Dana. I n “Bonitation of the World Ocean. 5 . Distribution and Biology of Commercial Invertebrates and Algae” (Ed. P. A. Moiseev), pp. 104-1 16. Pishchevaya promyshlenhost, Moscow. Pavlov, V. Ya. (1976a). Feeding habits of Euphausia triacantha Holt and Tattersall. Trudy V8esoyuznogoNauchno-issledovatel’skogoInstituta Morskogo Rybnogo Khozyaistva i Okeanografii (Trudy VNIRO), 110, 108-110. Pavlov, V. Ya. (197613). On the function of scaphocerites in euphausiids. T d y Vseaoyuznogo Nauchno-ksledovatel’skogo Instituta Morskogo Rybnogo Khozy aistva i Okeanografii (Trudy VNIRO), 110, 111-114. Paxton, J. R. (1967). Biological notes on southern California lantern fishes (Family Myctophidae).California Fish and Game, 53, 214-217. Pearcy, W. G . and Ambler, J. W. (1974). Food habits of deep-sea macrourid fishes off the Oregon coast. Deep-sea Research, 21, 745-759.

-

REFERENCES

583

Pearcy, W. G. and Hancock, D. (1978).Feeding habits of dover sole, Microstomwr pacijicwr ; rex sole, Olyptocephalh zachirwr ; slender sole, Lyopsetta exilis ; and Pacific sanddab, Cithurichthyssordidus in a region of diverse sediments and bathymetry off Oregon. Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 76, 641-651. Pearcy, W. G.and Mesecar, R. S. (1971).Scattering layers and vertical distribution of oceanic animals off Oregon. I n “Biological Sound Scattering in the Ocean” (Ed. G. B. Farquhar), pp. 381-394. Maury Center for Ocean Science, Department of the Navy, Washington, D.C. Pearcy, W. G., Theilacker, G. H. and Lasker, R. (1969).Oxygen consumption of Euphausia pacijca: the lack of a die1 rhythm or light-dark effect, with a comparison of experimental techniques. Limnology and Oceanography, 14, 219-223. Pearcy, W. G.,Krygier, E. E., Mesecar, R. and Ramsey, F. (1977). Vertical distribution and migration of oceanic micronekton off Oregon. Deep-sea Research, 24, 223-245. Pedenovi, C. and Croce, N. D. (1971).Biological results of sampling in the deep scattering layers of the western Mediterranean in autumn. Technical Report SACLANT ASW Research Centre, 189, 1-61. Pennitin, Yu. E. (1970). The consumption of krill by Antarctic fishes. I n “Antarctic Ecology” (Ed. M. W. Holdgate), Vol. 1, pp. 177-182. Academic Press, London and New York. Permitin, Yu. E. and Terverdieva, M. I. (1978).Feeding of Nototheniidae and Chaenichthyidae near the South Orkney Islands. Biologiya Morya ( Vladivostock), No. 2, 76-81. Pertzova, K.N.(1976).Larvae of Euphausiacea in the Antarctic. Trudy Instituta okeanologii, Akademiya nauk SSSR, 105, 147-170. Petersen, W. T. and Miller, C. B. (1975).Year-to-year variations in the planktology of the Oregon upwelling zone. Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 73, 642-653. Petersson, G. (1968).Studies on photophores in the Euphausiacea. Sarsia, 36,l-39. Petushko, G. L. (1969).The behaviour of krill in photo and electrical fields. Trudy Vsesoyuznogo Nauchno-issledovatel’skogoInstituta Morskogo Rybnogo Khozyaistwa i OkeanograJi, 66, 240-242. National Lending Library for Science and Technology, Boston Spa, Yorkshire. Translation RTS 5589. Pieper, R. E. (1977). Some comparisons between oceanographic measurements and high frequency scattering of underwater sound. I n “Oceanic Sound Scattering Prediction” (Eds N. R. Andersen and B. J. Zahuranec), Marine Science, Vol. 5,pp. 667-677. Plenum Press, New York and London. Pierce, R. W., Van der Veen, J. and Olcott, H. S. (1969).Proximate and lipid analyses of krill (Euphawria species) and red crab (Pleuroncodes planipes). Journal of Agricultural and Food Chemistry, 17, 367-369. Phhorn, A. T. (1976).Livingmarineresourcesof Newfoundland-Labrador :status and potential. Bulletin of the Fisheries Research Board of Canada, 194, 1-64. Pitt, T. K. (1973).Food of American plaice (H&ppoglossodesplatemodes) from the Grand Bank, Newfoundland. Journal of the Fisheries Research Board of Canada, 30, 1261-1273. Pommeranz, T. (1978).Attempts for quantitative studies in the distribution of adult krill Euphauaia superba Dana in the Scotia Sea. International Council for the Exploration of the Sea, Biological Oceanography Committee Paper, CM 1978/L:6, 1-17.

584

THE BIOLOGY OF EUPHATJSIIDS

Ponomarenko, I. Y. (1976). Food composition and quantitative indices of feeding of 0-group cod in the Barehts Sea. A n n a b biologiques, 31 (1974), 83-85. Ponomarenko, I. Y. (1978). Investigations on feeding and biological indices of 0-group cod in the Barents Sea. A n n a b biologiqws, 83 (1976), 81-82. Ponomareva, L. A. (1963). The Euphawri& of the North Pacifi, their Distribution and EcoEogy. Doklady Akademy Nauk, SSSR, 1-142. Israel Programme for Scientific Translation, Jerusalem, 1966. Ponomareva, L. A. (1969). Investigations on some tropical euphausiid species of the Indian Ocean. Marine Biology, 3, 81-86. Ponomareva, L. A. (1971). Circadian migrations and feeding rhythm of some Indmn Ocean euphausiid species. Oceanology, Moscow, 11, 226-231. Ponomareva, L. A. (1972). Quantitative distribution of euphausiids in the Indian Ocean. Oceanology, Moscow, 12, 576-581. Ponomareva, L. A. (1975). The Euphu&& of the Indian Ocean and Red Sea. Akademy Nauk, SSSR, Moscow, 83 pp. Ponomareva, L. A. (1976a). Data on the ecology of feeding in Euphausiacea from the Far Eastern Seas. I n “Bottom Fauna of the USSR Fringing Seas”, pp. 106-111. Academy Nauk, Moscow. Ponomareva, L. A. (197613). On the feeding habits of euphausiids in the northwest Pacific. Trudy Vseeoyuznogo Nauchno-Wskdovatel’skogoImtituta Morskogo Rybnogo Khozyaistva i Okeanografii (Trudy VNIRO), 110, 10P107. Ponomareva, L. A. (1976~).Euphausiids of the Sulu Sea and adjacent waters. oceanology, Moscow, 16, 5 1 4 4 1 6 . Ponomareva, L. A., Tsikhon-Lukanina, E. A. and Sorokin, Yu. I. (1971). On phytoplankton and bacteria consumption of tropical euphausiids. I n “Functioning of Pelagic Communities in Tropical Oceanic Regions”, pp. 152156. Institute of Oceanology, Academy Nauk, SSSR, Moscow. Postel, E. (1972). Le plancton, source de nourriture. La P2che Maritime, 1137, 1005-1011.

Pratap, M. M., Wafar, M. V. M., Haridas, P., Ntarayanan, B., Menon, P. G. and Sivadas, P. (1977). Comparative studies on the abundance of zooplankton in the surrounding sea and lagoons in the Lakshadweep. Indian Journal of Marine Sciences, 6, 138-141. Prygunkova, R. V. (1974). Certain peculiarities in the seasonal development of zooplankton in the Chupa Inlet of the White Sea. I n “Explorations of the Fauna of the Seas XI11 (XXI). Seasonal Phenomena in the Life of the White and Barents Seas”, pp. 4-55. Zoological Institute, Academy of Sciences of the USSR. Pugh, W. L. (1972). Collections of midwater organisms in the Cariaco Trench, Venezuela. Bulletin of Marine Science, 22, 592-600. Ragulin, A. G. (1969). Underwater observations on krill. Trudy Vseeoyuznogo Nauchno-issledova~l’skogoImtituta Morskogo Rybnogo Khozyaktva i Okeanografii (Trudy V N I R O ) , 66, 231-234. Ministry of Agriculture, Fisheries and Food, Lowestoft, Translation N.S. 92 (1970). Rakusa-Suszczewski, S. and MacWhinnie, M. A. (1976). Resistance to freezing by Antarctic fauna : supercooling and osmoregulation. Comparative BWchemistry and Physiology, 54A,291-300. Ramirez, F. (1971). Eufausidos de algunos sectores del Atlantic0 sudoccidental. PhyGs, Section A , Buenoa Airee, 30, 385-405. Ramirez, F. C. (1973). Eufausidos de la campana oceanografica “Walther Herwig” 1966. Physia, Section A , Buenos Aires, 32, 105-114.

REFERENCES

686

Ramirez, F. C. (1977). Planktonic indicators of Argentine shelf and adjacent areas. I n “Proceedings of the Symposium on Warm Water Zooplankton”, pp. 65-68. Special Publication of National Institute of Oceanography, Goa, India. Raymond, J. A. and De Vries, A. L. (1976). Bioluminescence in McMurdo Sound, Antarctica. Limnology and Oceanography, 21, 599-602. Raymont, J. E. G., Srinivasagam, R. T. and Raymont, J. K. B. (1969). Biochemical studies on marine zooplankton. IV. Investigation on Meganyctip h n e s norvegica (M. Sars). Deep-sea Research, 16, 141-156. Raymont, J. E. G., Srinivasagam, R. T. and Raymont, J. K. B. (1971a). Biochemical studies on marine zooplankton. VIII. Further investigations on Meganyctiphanes norvegica (M. Sers). Deep-sea Research, 18, 1167-1178. Raymont, J. E. G., Srinivctsagam, R. T. and Raymont, J. K. B. (1971b). Biochemical studies on marine zooplankton. I X . The biochemioal composit,ionof Euphausia superba. Journal of the Marine Biological Association of the United Kingdom, 51, 581-588. Razouls, S. and Thiriot, A. (1968). Le macroplankton de la region de Banyulssur-Mer (Golfe de Lion). Vie et Milieu, 19 (lB), 133-184. Reeve, M. R., Grice, G. D., Gibson, V. R., Walter, M. A., Darcy, K. and Ikeda, T. (1976). A controlled environmental pollution experiment (CEPEX) and its usefulness in the study of larger marine zooplankton under toxic stress. I n “Effects of Pollutants on Marine Organisms” (Ed. A. P. Lockwood), pp. 145162. Cambridge University Press, Cambridge. Reeve, M. R., Gamble, J. C. and Walter, M. A. (1977).Experimental observations on the effects of copper on copepods and other zooplankton: controlled ecosystem pollution experiment. Bulletin of Marine Science, 27, 92-104. Reid, J. L. (1977). Some thoughts on the dependence of sound speed and the scattering layers upon ocean circulation. I n “Oceanic Sound Scattering Predictions” (Eds N. R. Andersen and B. H. Zahuranec), Marine Science, Vol. 5, pp. 15-64. Plenum Press, New York and London. Reuben, S. (1968). On the occurrence of E u p h u s i a distinguenda Hansen in the northwestern Bay of Bengal. Current Science, 37, 703-704. Rice, A. L. (1967). The orientation of the pressure responses of some marine Crustecea. I n “Symposium on Crustacea”, Vol. 3, pp. 112P1131. Marine Biological Association of India. Robertson, D. A. and Roberts, P. E . (1978). Mesopelagic faunal transition across the subtropical convergence east of New Zealand. New Zeahnd Journal of Marine and Freshwater Research, 12, 295-312. Roe, H. S. J. (1974). Observations on the diurnal vertical migrations of a n oceanic animal community. Marine Biology, 28, 99-1 13. Roger, C. (1968a). Evaluation d’une population planctonique. Marine Biology, 2, 66-70. Roger, C. (1968b). Note sur la rBcolte des euphausiachs p6lagiques. Cahiers ORSTOM-Ocdanographie, 6 , 3-1 1. Roger, C. (1968~). Premiere sBrie d’observations sur les euphausiaces du Pacifique ouest Bquatorial et tropical sud. Cahiers ORSTOM-Ochnographique, 6, 83-96. Roger, C. (1968d). Euphausiac6s des Taumotous. Cahiers 0RSTOM-Ocdanographie, 6, 31-38. Roger, C. (1971). Distribution verticale des euphausiac6s (Crustacbs) dans les courant8 Bquatoriaux de l’Oc6an Pacifique. Marine Biology, 10, 134-144.

686

TEE BIOLOGY OF EWPHAUSXIDS

Roger, C. (1973a). Recherches sur la situation trophiques d’un groupe d’orgmismes pblagiques (Euphausiacea). ‘I.Niveaux trophiques des esphces. Marine Biology, 18, 312-316. Roger, C. (197313).Recherches sur la situation trophique d’un groupe d’organismes pBlagques (Euphauaiacea). 11. Comportements nutritionnels. Marine Biology, 18, 317-320. Roger, C. (1973~). Recherches sur la situation trophique ci’un group d’organismes p6lagiques (Euphausiacea). 111. Potential alimentaire du groupe. Marine Biology, 18, 321-326. Roger, C. (1973d). Recherches sur la situation trophique d’un groupe d’organisme pBlagiques (Euphausiacea). IV. Relations avec lea autres BlBments du micronecton. Marine Biology, 19,54-60. Roger, C. (1973e).Recherches sur la situation trophique d’un groupe d’orgenismes pBlagiques (Euphausiacea). V. Relations avec lea thons. Marine Biology, 19, 61-65. Roger, C. (1973f).Recherches sup la situation trophique d’un groupe d’organismes pBlagiques (Euphausiacea).VI. Conclusions sur le r61e des euphausiacbs dans les circuits trophiques de l’Oc6an Pacifique intertropical. Marine Biology, 19, 66-68. Roger, C. (19738).Biological investigations of some important species of Euphausiacea (Crustacea) from the equatorial and south tropical Pacific. I n “Oceanography of the South Pacific” (Ed. R. Fraser), pp. 449-456. New Zealand National Commission for UNESCO, Wellington. Roger, C. (1974a). Influence de la phase et de 1’6clairement lunaire sur lea r6partitions verticales nocturnes superficielles de crustacBs macroplanctoniques (Euphausiacea). Cahiers ORSTOM-Ocdanographie, 12, 159-171. Roger, C. (197410). RBpartitions bathymBtriques et migrations verticales des euphausiac6s (crustac6s) dans lea zones de p6che au thon du Pacifique sudtropical. Cahiers ORSTOM-Ochnographie, 12, 221-239. Roger, C. (1974~).Lea euphausiaces du Pacifique Bquatorial et sud-tropical. ZoogBographie, Bcologie, biologie, et situation trophique. Memirea 0RSTOM -Ocdanographie, No. 71, 1-265. Roger, C. (1975). Rythmes nutritionnels et organisation trophique d’une population de crustac6s p6lagiques (Euphausiacea). Marine Biology, 32, 365-378. Roger, C. (1976). Fecundity of tropical euphausiids from the central and western Pacific Ocean. Crustaceam, 31, 103-105. Roger, C. (1977). The use of a group of macroplanktonic organisms (euphausiid crustaceans) in the study of warm water pelagic food webs. I n “Proceedings of the Symposium on Warm Water Zooplankton”, pp. 309-318. Special Bulletin of National Institute of Oceanography, Goa, India. Roger, C. (19788,). Bioecological sheets on tropical Pacific euphausiids. L’Ofice de la Recherche Scienti$que et Technique Outre-Mer, Initiations-Documentations Techniques, No. 36, 1-81. Roger, C. (197813). Azote et phosphore chez un crustace macroplanctonique, Meganyctiphanes norwegica (M. Sara) (Euphausiacea) : excretion min6rale et constitution. Journal of Experimental Marine Biology and Ecology, 33,57-83. Roger, C. and Grandperrin, R. (1976). Pelagic food webs in the tropical Pacific. fimnology and Oceanography, 21, 731-735. Roger, C.and Wauthy, B. (1968). Sur une technique de determination de groupes de tailles applicable 8, 1’6tude de certains organiemes planctoniques. Journal du Conseil, 37, 162-169.

REFERENCES

587

Rorvik, C. J., Jonsson, J., Mathisen, 0. A. and Jonsgard, A. (1976). Fin whales, Balaenoptera physalus (L.), off the west coast of Iceland, segregation by length and exploitation. Rit Fiskideildar, 5, 1-30. Roschke, N. (1976). Chemische Analysen von Krill- und Fischprodukten der deutschen Antarktis-Expedition 1975/76 : Methoden und erste Ergebnisse. Informations Fischwirtschaft, 23, 1 3 6 1 3 9 . Roschke, N. (1977). Haltbarkeit von Antarktischen Fischen und Krillprodukten. Informations Fischwirtschaft, 24, 121-123. Roschke, N. (1978). Variabilitiit des Fett- und Proteingehalts beim Krill (Euphausia superba Dana). Meeresforschung, Reports o n Marine Research, 26, 2 17-22 1. Roschke, N. and Schreiber, W. (1977). Analytik von Krill, Krillprodukten und antarktischen Fische. Archiv fiir Fischereiwissenschaft, 28, 135-141. Rottman, M. L. (1978). Ecology of recurrent groups of pteropods, euphausiids, and chaetognaths in the Gulf of Thailand and the South China Sea. Marine Biology, 48, 63-78. Rudjakov, J. A. (1970). The possible causes of die1 vertical migrations of planktonic animals. Marine Biology, 6,98-105. Rudjakov, J. A. and Tseitlin, V. B. (1976). Characteristics of the diurnal vertical migrations of the pelagic fauna in the Canary Islands area. Okeanologija, MOSCOW, 16, 329-335. Ruud, J. T. (1932). On the biology of southern Euphausiidae. Hvzllradete Skrifter, (2), 1-105. Sakasai, N., Hayashi, K., Nakajima, G., Saito, N. and Shiba, M. (1978). A convenient assay method for protease of the Antarctic krill using paper disks. Bulletin of the Japanese Society of Scienti$c Fisheriee, 44, 673-675. Sameoto, D. D. (1972). The feasibility of mapping euphausiid concentrations in the Gulf of St. Lawrence using high frequency sonar. Fishery Research Board of Canada, Technical Report, No. 350, 1-30. Sameoto, D. D. (1976a). Distribution of sound scattering layers caused by euphausiids and their relationship to chlorophyll a concentrations in the Gulf of St. Lawrence estuary. Journal of the Fisheries Reaearch Board of Canada, 33, 681-687. Sameoto, D. D. (1976b). Respiration rates, energy budgets and molting frequencies of three species of euphausiids found in the Gulf of St. Lawrence. Journal of the Fisheries Research Board of Canada, 33, 2568-2576. Sameoto, D. D. (1978). Zooplankton sample variation on the Scotian Shelf. Journal of the Fisheries Research Board of Canada, 35, 1207-1222. Sameoto, D. D., Darrow, D. C. and Guildford, S. (1975). DDT residues in the upper estuary of the Gulf of St. Lawrence. Journal of the Fisheries Raearch Board of Canada, 32, 310-314. Samyshev, E. Z. and Schetinkin, S. V. (1973). Feeding patterns of some species of Myctophidae and Maurolicus muelleri caught in the sound-dispersing layers in the north-western African area. Annales biobgipw, 28 (1971), 212-2 15. Sargent, J. R. (1976). The structure, metabolism and function of lipids in marine organisms. I n “Biochemical and Biophysical Perspectives in Marine Biology” (Eds D. C. Malins and J. R. Sargent), Vol. 3, pp. 150-212. Academic Press, New York and London. Sargent, J. R . and Lee, R. F. (1975). Biosynthesis of lipids in zooplankton from Saanich M e t , British Columbia, Canada. Marine Biology, 31, 15-23.

688

THE BIOLOGY OF ETJT’HATJSIIDS

Sarhage, D. (1978). Zur Llingen-Gewichts-Beziehungbeim Antarktischen Krill (Euphausia superba). Meeresforschung, Reports on Marine Research, 26, 47-49. Sarhage, D. and Steinberg, R. (1975). Dem Krill auf der Spur Expedition in Antarktische Gewiiaser. U m c h a u , 20, 627-631. Sarhage, D., Schreiber, W., Steinberg, R. and Hempel, G. (1978). Antarctic expedition 197511976 of West Germany. Archiv fur Fischereim‘ssenschaft, 29, Suppl. 1, 1-96. Sers, G. 0. (1885). Report on the Schizopoda collected by H.M.S. “Challenger” during the years 1873-1876. The Voyage of H.M.S. “Chalbnger”, 13 (37), 1-228. Sasaki, Y., Inoue, K. and Matsuike, K. (1968). Distribution of Euphausia superba in the Atlantic Ocean from view points of the fishing operation. Journal of the Tokyo University of Fisheries, 9, 129-134. Sawamoto, S. (1976). New records of Euphawria gibba and Euphawria sibogae, (Euphausiacea)from the western part of the tropical and subtropical North Pacific Ocean. Journal of the Faculty of M a r i m Science and Technology, Tokai University, 9, 65-71. Schnack, D. (1978). Comments on sampling tests made with Beyer’s epibenthic closing net. Meeresforschung, Reports on Marine Research, 26, 11-14. Sebastian, M. J. (1966). Euphausiacea from Indian Seas: systematics and general considerations. I n “Symposium on Cmstacea”, Vol. 1, pp. 233-254. Marine Biological Association of India. Sebastian, M. J. (1970). On two isopod parasites of Indian euphausiids. Journal of Natural History, 4, 153-158. Sedberry, G. R. and Musick, J. A. (1978). Feeding strategies of some demersal fishes of the continental slope and rise off the mid-Atlantic coast of the USA. Marine Biology, 44, 357-375. Seki, N., Ozawa, R. and Arai, K. (1975). Studies on muscle proteins of Antarctic krill. Bulletin of the Japanese Society of Scienti$c Pisherim, 41, 1287-1292. Seki, N., Sakaya, H. and Onozawa, T. (1977). Studies on proteases from Antarctic krill. Bulletin of the Japanese Society of Scientijk Fisheries, 43, 955-962. Selvakumaran, M., Hodson, R. and Menzies, R. J. (1974). The effects of hydrostatic pressure on living aquatic ‘organisms. VII. Size and pressure effects. Internationale Revue der gesamten Hydrobiologie u. Hydrographie, 69, 213-218. Semenov, V. N. (1969). Aquarium observations on the behaviour of krill. T d y Vsesoyuznogo Nauchno-idedovatel’skogo Instituta Morskogo Rybnogo Khozyaiatva i Okeanografii (Trudy V N I R O ) , 66, 235-239. Ministry of Agriculture, Fisheries and Food, Lowestoft, Translation No. 93 (1970). Sergeant, D. E. (1973). Feeding, growth, and productivity of northwest Atlantic harp seals (Pagophilw groenladicua). Journal of the Fisheries Research Board of Canada, 30, 17-29. Shannon, L. V. (1972). Marine alpha-activity of southern Africa. I. Gross alphaactivity, radiation dose, alpha-spectrum and variations in alpha-activity of marine life. Investigational Report of the Division of Sea Fisheries, South Africa, 98, 1-80. Sheard, K. ( 1953). Taxonomy, distribution and development of the Euphausiacea (Crustacea). Report of the British and New Zealand Antarctic Research Expedition, Ser. B (Zoology and Botany), 8, 1-72. Sheard, K. (1965). Species groups in the zooplankton of eastern Australian slope waters, 1938-41. Awtralian Journal of Marine and Freshwater Research, 16, 21 9-264.

REFERENOES

689

Sheard, K. (1967). Temperature and chlorinity distributions, and species associations of some subtropical euphausiid and hyperiid Crustacea. I n “Symposium on Crustacea”, Vol. 3, pp. 976-986. Marine Biological Association of India. Shevtsov, V. V. and Makarov, R. R. (1969). On the biology of the Antarctic krill. Trudy Vsseaoyuznogo Nauchno-~88ledovatel’skogo Instituta Morskogo Rybnogo Khozyaistva i Okeanografii (Trudy VNIRO), 66, 177-206. Ministry of Agriculture, Fisheries and Food, Loweatoft, Translation No. 91 (1970). Shih, C.-T. and Laubitz, D. R. (1978). Zooplankton distribution in the eastern Beaufort Sea and the Northwest Passage. Astarte, J o u d of Arctic Biology, 11, 45-54. Shimazu, T. (1975). Some cestode and acanthocephalan larvae from euphausiid crustaceans collected in the northern North Pacific Ocean. Bulletin of t7w Japanese Society of Scientifi Fieheria, 41, 813-821. Shimazu, T. and Kagei, N. (1978). A metacercaria of.the genus P a r o d r e m a (Trematoda: Syncoeliidae)parasitic in Euphausia pacifica (Crustacea:Euphausiidae) from the East China Sea. Zoological Magazine, Tokyo, 87, 168-161. Shimazu, T. and Oshima, T. (1972). Some larval nematodes from euphausiid crustaceans. I n “Biological Oceanography of the Northern North Pacific Ocean” (Eds A. Y. Takenouti et al.),pp. 403-409. Idemitm Shoten, Tokyo. Shomura, R. S. and Nakamura, E. L. (1969). Variations in marine zooplankton from a single locality in Hawaiian waters. Piehey Bulletin, National Oceanic Atmospheric Adminietratwn of the United Stater,, 68, 87-100. Shust, K. V. (1969). Visual observations on Antarctic krill made on board the factory ship “Akademik Knipovich”. Trudy Vser,oyuznogo Nauchm-Gel&vatel’skogo Inatituta Morebgo Rybnogo Khozyaietva i OkeanograJG (Trudy VNIRO),66,223-230. National Lending Library for Science and Technology, Boston Spa, Yorkshire, Translation RTS 5588 (1970). Shvetsova, G. M. (1977). On biology of Euphawiacea of the south-western part of the Sea of Okhotsk. Qidrobiologicbkii Z h u d . 13, 2 6 2 8 . Sidhu, G. S., Montgomery, W. A., Holloway, G . L., Johnson, A. R. and Walker, D. M, (1970). Biochemical composition and nutritive value of krill (EuphazLeia auperba Dana). Journal of the Science of Food and Agriculture, 11, 293-296. Silas, E. G. and Mathew, K. J. (1967). Stylocheiron indicw, a new euphausiid (Crustacea: Euphausiacea) from Indian Seas. Current Science, 36, 169-172. Silas, E. G. and Mathew, K. J. (1977). A critique to the study of larval development in Euphausiacea. I n “Proceedings of the Symposium on Warm Water Zooplankton”, pp. 57 1-582. Special Publication of National Institute of Oceanography, Goa, India. Sipos, V. (1977). Distribution of euphausiids in the Adriatic Sea in autumn 1974 and spring 1975. Rapport8 et procbverbam o h rduniona. C m i e e i o n interd W e pour l’exploratwn scientifique de la rner Miditerrane’e,24, 123-124. Skjoldal, H. R. and B&mstedt, U. (1976). Studies on the deep-water pelagic community of Korsfjorden, western Norway. Adenosine phosphates and nucleic acids in Meganyctiphaner, norvegica (Euphawiacea) in relation to the life cycle. Sarsia, 61, 1-14. Skjoldal, H. R. and BBmstedt, U. (1977). Ecobiochemical studies on the deepwater pelagic community of Korsfjorden, western Norway. Adenine nucleotides in zooplankton. Marine Biology, 42, 197-211. Sluiters, J. F. (1973). Study on the life cycle of Anieakie sp. Parasitology, 67, p. xxviii.

690

THE BIOLOGY OF EUPHAUSIIDS

Small, L. F. (1969). Experimental studies on the transfer of 06Zn in high concentration by euphausiids. Jotirnal of Experimental Marine Biology and Ecology, 3, 10G123.

Small, L. F. and Fowler, S. W. (1973). Turnover and vertical transport of zinc by the euphausiid Meganyctiphanes norvegica in the Ligurian Sea. Marine Biology, 18, 284-290. Small, L. F., Fowler, S. W. and KeZkeS, S. (1973). Flux of zinc through a macroplanktonic crustacean. I n “Radioactive contamination of the Marine Environment”, pp. 437-452. International Atomic Energy Agency, Vienna. Smeyda, T. J. (1969). Some measurements of the sinking rate of fecal pellets. Limnology and Oceanography, 14, 621-625. Smiles, M. C. and Pearcy, W. G. (1971). Size structure and growth rate of EuphazLsia pacifica o f f the Oregon coast. Fishery Bulletin, National Oceanic Atmospheric Administration of the Unikd States, 69, 79-86. Smith, J. W. (1971). Thyaanohsa inermb and T . long~mudatu(Euphausiidae) as first intermediate hosts of Anisakk sp. (Nematoda:Ilscaridata) in the northern North Sea, to the north of Scotland and Faroe. Nature, London, 234, p. 478. Soboleva, M. S. (1974). Distribution and abundance of euphausiids in the autumnwinter period of 1971-72 in the southern Barents Sea. Annales biologiques, 29, (1972), 40-41.

Soboleva, M. S. (1975). The distribution and abundance of euphausiids in the Barents Sea in autumn and winter 1972-73. Annales biologiques, 30, (1973), 58-60.

Southward, A. J. (1962). The distribution of some plankton animals in the English Channel and approaches. 11. Surveys with the Gulf I11 high-speed sampler, 1958-60. Journal of the Marine Biological Association of the United Kingdom, 42, 275-375. Srinivasagam, R. T., Raymont, J. E. G., Moodie, C. F. and Raymont, J. K. B. (1971). Biochemical studies on marine zooplankton. X. The amino acid composition of Euphawia superba, Meganyctiphanes norvegica and Neomyeis integer. Journal of the Marine Biological Association of the United Kingdom, 51, 917-925.

Steinberg, a. and Mentjes, T.(1978). Zum Stand der Ortung und Fangtechnik in der antarktischen Krillfischerei. Informations Fischwirtschaft, 25, 25-27. Suyama, M., Nakajima, K. and Nonaka, J. (1965). Studies on the protein and non-protein nitrogenous constituents of EuphaauSia. Bulletin of the Japanese Society of Scientific Fisheries, 31, 302-306. Suzuki, T. (1973). Ecological studies on the jack mackerel, Trachurus japonicw, (Temminck and Schlegel). 111. Amount of food consumption for growth of young fish. Bulletin of the Japan Sea Regional Fisheries Research Laboratory, 24, 67-76.

Suzuki, T. end Kanna, K. (1977). Denaturation of muscle protein in krill and shrimp during frozen storage. Bulktin of the Tokai Regional Sea Fisheries Laboratory, No. 91, 67-72. Swift, E., Biggley, W. H. and Napora, T. A. (1977). T h e bioluminescenceemission spectra of Pyrosoma atlanticum, P. spinosum (Tuniceta), Eupha& tenera (Crustacea)and Gonostomu (Pisces).Journal of the Marine Biological Association of the United Kingdom, 57, 817-823. Takagi, M., Iida, A. and Oka, S. (1975). PCB contents in several species of zooplankton collected in the North Pacific Ocean. BJletin of the Japanese Society of Scientific Fbheries, 41, 561-565.

REFERENCES

591

Takahashi, H. and Yamada, M. (1976). Lipid,composition of seven species of crustacean plankton. Bulletin of the Japanese Society of Scientific Fisheries, 42, 769-176.

Takahashi, M. and Ikeda, T. (1975). Excretion of ammonia and inorganic phosphorus by Euphausia pacifica and Metridia pacificu at different concentrations of phytoplankton. Journal of the Fisheries Research Board of Canada, 32, 2189-2195.

Talbot, M. S. (1974). Distribution of euphausiid crustaceans from the Agulhas Current. Zoologica Africana, 9, 93-145. Taniguchi, A. (1967). Abnormal male specimen of Nematoacelia tenella (Euphausiacea) having dorsal keels. Information Bulletin of Planktology, Japan, No. 14, 69-60.

Taniguchi, A. (1969). Mysidacea and Euphausiacea collected in the south-east of Hokkaido, Japan. Bulletin of the Faculty of Fiaheries, Hokkaido University, 20, 43-59.

Taniguchi, A. (1976). Mysids and euphausiids in the eastern Indian Ocean with particular reference to invasion of species from the Banda Sea. Journal of the Marine BiologicaE Aaaociation of India, 6, 349-357. Tarverdiyeva, M. I. (1972). Daily food consumption and feeding pattern of the Georgian cod (Notothenia roaai nzamnorata Fischer) and the Patagonian toothfish (Baaostichua eleginoidaP Smitt) (Fam. Nototheniidae) in the South Georgia area. Voproay ikhtiologii (English edition), 12, 684-692. Tattersall, W. M. (1924). Crustacea. VIII. Euphausiacea. Bm’tiah Antarctic “Terra Nova” Expedition, 1910, Zoology, 8, 1-36. Taylor, P. M. (1976). Bibliography on krill as a human food source, 1-9. Ministry of Agriculture, Fisheries and Food, Marine Laboratory, Tony, Aberdeen. Tett, P. B. (1969). The effects of temperature upon the flash-stimulated luminescence of the euphausiid Thyaanoaaa raachii. Journal of the Marine Biological Aaaociation of the United Kingdom, 49, 245-258. Tett, P. B. (1972). An annual cycle of flash induced luminescence in the euphausiid Thyaanoaaa raachii. Marine Biology, 12, 207-218. Tett, P. B. and Kelly, M. G. (1973). Marine bioluminescence. Oceanography and Marine Biology, a n Annual Review, 11, 89-173. Tkachenko, V. N. and Patin, S. A. (1971). Artificial radioactivity of some hydrobionts of the Southern Ocean (Scotia Sea). Oceanology, Moscow, 11, 694-598.

Tokunaga, T., Iida, H. and Nakamura, K. (1977). Formation of dimethyl sulfide in Antarctic krill, Euphauaia auperba. Bulletin of the Japanese Society of Scientijk Fiaheries, 43, 1209-1217. Tolkach, V. V. and Gromov, V. V. (1975). Absorption of Sr@o by krill. Okeanologija, Moacow, 15, 639-642. Tolkach, V. V. and Gromov, V. V. (1976).Yttrium-90uptake by krill.OkeanoZogija Moscow, 16, 1015-1019. Tsuji, F. I., Haneda, Y., Lynch, It. V. and Sugiyama, N. (1971). Luminescence cross-reactions of Porichthya luciferin and theories on the origin of luciferin in some shallow-water fishes. Comparative Bwchemiatry and Physiology, 40A, 163-179.

Tsuyuki, H. and Itoh, S. (1976). Fatty acid component of lipid of EuphaUaia auperba. Scientific Reports of the Whales Research Institute Tokyo, No. 28, 107-174.

592

THE BIOLOGY OF EUPHAWSIIDS

Tupolev, V. M. (1969). The catching of the Antarctic krill on an experimentalcommercial basis from the SR ‘TR’s “Orekhovo” and “Obdorsk”. Trudy Vsesvyuznogo Nauchno-issledovatel’skogoInstituta Morskogo Rybnogo Khozyaistva i Okeanogra$i (Trudy V N I R O ) ,66, 284-294. National Lending Library for Science and Technology, Boston Spa, Yorkshire, Translation RTS 659’3 (1970). Tyler, A. V. (1972). Food resource division among northern marine demerml fishes. Journal of the Fisheries Research Board of Canada, 29, 997-1003. Tyler, A. V. (1973). Caloric values of some north Atlantic invertebrates. Marine Biology, 19, 258-261. Tyler, H. R. and Pearcy, W. G. (1975). The feeding habits of three species of lantern fishes (family Myctophidae) off Oregon, U. S.A. Marine Biology, 32, 7-11. Uspenskaya, A. V. (1963). “Parasite-Fauna of Benthic Crustaceans in the Barents Sea”. Murmanskii Morskoii Biologischeskii Institut. Akademia Nauk, SSSR, Moscow. Vader, W. (1973). A bibliography of the Ellobiopsidae, 1959-1971, with a list of Thalassomyces species and their hosts. Sarsia, 52, 175-180. Valentine, J. W. and Ayah, F. J. (1976). Genetic variability in krill. Proceedings of the National Academy of Science, United States, 73, 658-660. Vanderploeg, H. A. (1979). Dynamics of zinc-65 specific activity and total zinc in benthic fishes on the outer continental shelf off central Oregon. Marine Biology, 52, 2 5 9-2 72. Van der Veen, J., Medwadowski, B. and Olcott, H. S. (1971). The lipids of krill (Euphausia species) and red crab (Pleuroncodes planipes). Lipids, 6, 481-485. Vinogradov, M. E. (1970). “Vertical distribution of the oceanic zooplankton”, pp. 1-339. Israel Pr0gra.m for Scientific Translations, Jerusalem. Vinogradov, M. E. and Parin, N. V. (1973). Some features of the vertical distribution of macroplankton in the tropical Pacific. Okeanologija, Moscow, 13, 137-148. Vinogradov, Z. A. and Kandiuk, R. P. (1967). Sterols, provitamins D and cholesterol in marine organisms. I n “Biochemistry of Marine Organisms”, pp. 61-69. Ukranian Academy of Sciences, SSR. Vives, F. (1968). Sur 10s malacostrac6s planotoniques des mers Tyrrhhienne et Catalane. Rapport et procb-verbaux des rhunions. Commission internationale pour l’exploration scientijique de la mer Mhditerranhe, 19,459-461. Vlymen, W. J. (1970). Energy expenditure of swimming copepods. Limnology and Oceanography, 15, 348-356. Voronina, N. M. (1968). The distribution of zooplankton in the Southern Ocean and its dependence on the circulation of the water. Sarsia, 34, 277-284. Voronina, N. M. (1974). An attempt at a functional analysis of the distributional range of Euphausia superba. Marine Biology, 24, 347-352. Voronina, N. M. and Naumov, A. G. (1968). Quantitative distribution and composition of mesoplankton in the Southern Ocean. Okeanologija, Moscow, 8, 1059-1065. Wang, R. (1965). On the larval stages of Pseudeuphausia sinica Wang and Chen (Euphausiacea). Oceanologiaet Limnologiu Sinica, 7,35-53. Watanabe, T., Sugii, K., Yuguchi, H. and Kinumaki, T. (1976). Studies on the utilization of Antarctic krill, Euphausia superba Dana-11. Analyses of nutritive components. Bulletin of the Tokai Regional Fisheries Reeearch Laboratory, No. 85, 13-30.

REFERENCES

593

Weigmann, R. (1970). Zur Okologie und Erniihrungsbiologie der Euphausiaceen (Crustacea) i m Arabischen Meer. “Meteor”-Forschungsergebnisse, D, No. 5, 11-52. Weigmann, R . (1971). Eine isolierte population von Psedeuphausia latifrom (Crustacea: Euphausiacea) im Persischen Golf. Marine Biology, 8, 351-365. Weigmann, R . (19744. Untersuchungen zur Zoogeographie der Euphausiaceen (Crustacea) des Roten Meeres. Helgolander wiesenschajtliche Meerecreuntersuchungen, 26, 225-237. W e i g m m , R. (1974b). Untersuchungen zum Vorkommen der Euphausiaceen (Crustacea) im Bereich der Groben Meteorbank. “Meteor”-Forschungsergebnisse, D, No. 17, 17-32. Weigmann-Haass, R. (1 976). Verbreitung und Verteilung der Euphausiacea (Crustacea) Un Auftriebsgebiet vor NW-Afrika. “Meteor”- Reise 19(1970) und 26(1972). “Meteor”- Forschwnqsergebnkse, D, No. 23, 62-72. Wiegmann-Haass, R . (1977). Dei calyptopis- und furcilia-stadien von Euphausia hanaeni (Crustacea : Euphausiacea). HelgollEnder wissenschaftlicheMeeresuntersuchungen, 29, 316-327. Wiborg, K. F. (1968). Atlantic euphausiids in the fjords of western Norway. Sarsia, 33, 35-42. Wiborg, K. F. (1971). Investigations on ouphausiids in some fjords on the west coast of Norway in 196G1969. Fiskeridirektoratets skrifter, Serie Havundersokelser, 16, 10-35. Wiebe, P. H. (1970). Small-scale spatial distribution in oceanic zooplankton. Limnology and Oceanography, 15, 205-217. Wiebe, P. H. (1972). A field investigation of the relationships between length of tow, size of net and sampling error. Journal du Comeil, 34, 268-275. Wiebe, P. H. and Boyd, S. H. (1978). Limits of Nenzatoscelis megalops in the northwestern Atlantic in relation to Gulf Stream cold core rings. I. Horizontal and vertical distributions. Journal of Marine Research, 36, 119-142. Wiebe, P. H. and D’Abramo, L. (1972). Distribution of euphausiid assemblages in the Mediterraean Sea. Marine Biology, 15, 139-149. Williamson, D. I. (1969). Names of larvae in the Decapoda and Euphausiacea. Crustuceana, 16, 210-213. Win, K. (1977). A species list of the zooplankton from the coastal waters of Burma. I n “Proceedings of the Symposium on Warm Water Zooplankton”, pp. 93-99. Special Publication of National Institute of Oceanography, Goa, India. Wing, B. L. (1974). Kinds and abundance of zooplmkton collected by the USCG icebreaker Gbcier in the eastern Chukchi Sea, September-October, 1970. Nutionab Oceanic Atmospheric A d ~ i n i s t r ~ t i oofn the United States, Technical Report, NlMFS SSRF-679, 1-18. Wing, B. L. (1975). New records of Ellobiopsidae (Protista incertae sedis) from the North Pacific with a description of Thaluasomyces albatrossi n.sp., a parasite of the mysid Stylomysis major. Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 7 3 , 169-185. Wing, B. L. and Barr, N. (1977). Midwater invertebrates from the southeastern Chukchi Sea: species and abundance in catches incidental to midwater trawling survey of fishes, September-October, 1970. National Oceanic Atmospheric Administration of the United States, Technical Report, NMFS SSRF-710, 1-43.

594

THE BIOLOQY OF EUPHAUSIIDS

Wolfe, D. C. and Webb, B. F. (1975). Slender tuna (AUothunnuafallai Serventy), first record of bulk catches, Tasdania, 1974. Australian Journal of Mwine and Freshwater Research, 26, 213-221. Wooten, R. (1978). The occurrence of larval anisakid nematodes in small gadoids from Scottish waters. Journal of the Marine Biological Association of the United Kingdom, 58, 347-366. Wroblewski, J. S. (1977). Vertically migrating herbivorous plankton-their possible role in the creation of small scale phytoplankton patchiness in the ocean. I n “Oceanic Sound Scattering Prediction” (Eds N. R. Andemen and B. J. Zahuranec), Marine Science, Vol. 5, pp. 817-847. Plenum Press, New York and London. Yamaguchi, I. (1975). Oplophorm oxyluciferin and a model luciferin compound biologically active with Oplophorua luciferase. The Biochemical J o u m l , 151, 9-16.

Yamamoto, T., Yamoaka, T., Fujita, T. and Isoda, C. (1973). Boron content of marine plankton. Record8 of Oceanographic Worka, Japan, 12, 13-21. Yanase, M. (1971). Chemical composition of Eqhausia auperba and its utilization as condensed solubles for human food. Bulletin of the Tokai Regional Fkheriea Research Laboratory, 65, 69-66. Yanase, M. (19744. Chemical composition of Antarctic krill Euphausia auperba by raw freezing and pre-cooked freezing. Bulletin of the Tokai Regional Fisheries Research Laboratory, 77, 97-102. Yanase, M. (197413). Modification of Russian method for separating heat coagulated protein from Antarctic krill. Bulletin of the Tokai Regional Fkheriea Research Laboratory, 78, 79-84. Yanase, M. (1975). Chemical composition of the exoskeleton of Antarctic krill. Bulletin of the Tokai Regional FkherieS Reaearch Laboratory, 83, 1-6.

Youngbluth, M. J. (1975). The vertical distribution and diel migration of euphausiids in the central waters of the eastern south Pacific. Deep-sea Research, 22, 519-636. Youngbluth, M. J. (1976). Vertical distribution and diel migration of euphausiids in the central region of the California Current. Fishery Bulletin, National Oceanic Atmmapheric Administration of the United States, 74, 925-936.

Zelickman, E. A. (1968). Some features of evolution of the family Euphausiidae (Crustacea, Euphausiacea) in neritic and oceanic areas. Zoologicheskii Z h u d , Moacow, 47, 1314-1327. Zelickman, E. A. and Golovkin, A. N. (1972). Composition, structure and productivity of neritic plankton communities near the bird colonies of the northern shores of Novaya Zemlya. Marine Biology, 17, 265-274. Zenkovich, B. A. (1970). Whales and plankton in Antarctic waters. I n “Antarctic Ecology” (Ed. M. W. Holdgate), Vol. 1, pp. 183-186. Academic Press, London and New York. Zerbib, C. (1967). Premiere observation de la glande androghe chez un crustac6 syncaride: A w p i d e s ti-mnamiae Thomson et chez un crustac6 eucaride: Meganyctiphanea m e g i c a Sam. Compte r e d u de l’deademie des Sciencea, Paris, 265, Ser. D, 416-418. Zhuravlev, V. M. (1976a). Life span of euphausiids from the Sea of Okhotsk. Trudy Vaeaoyuznogo Nawhno-kaledovatel’akogo Institutu Morakogo Rybnogo Khozyaktva i OkeunograJii (Trudy VNIRO), 110, 97-103.

REFERENCES

595

Zhuravlev, V. M. (197613).Diurnal fluctuations in the abundance of euphausiids in the 0-50m layer of the Okhotsk Sea. Trudy V7seaoyuznogo Nauchnohaledovatel'akogo Inslitutcc Morekogo Rybnogo Khozyahtva i OkeanograJii ( T d y V N I R O ) , 110, 77-80. Zhuravlev, V. M. (1977). Species composition and distribution of euphausiids in the Sea of Okhotsk. Okeanologija, Moscow, 17, 127-131. Zimmer, C. (1914). Die Schizopoden der Deutschen Sudpolar-Expedition, 19011903. Deutschm Sdpolar-Expedition, 15, Zoologie (7), 377-445.

ADDENDUM TO BIOLOGY OF EUPHAUSIIDS Additional papers have been published since completion of the original manuscript. A significant portion of these papers record work done on the Antarctic Euphausia superba. Many of these projects have been stimulated by the impending and current commercial exploitation of this species. A multi-national programme of Biological Investigations of Marine Antarctic Stocks and Systems-BIOMASS-is being planned at present (BIOMASS, 1977, 1978; El-Sayed, 1978; El-Sayed and McWhinnie, 1979; Tranter and Smith, 1979). I n this connection, a further bibliography on Antarctic krill and fishery resources has been published by Koniecka et al. (1977). The additional information is summarized here under the chapter numbers and headings of the main paper to facilitate easy crossreference. 2. The species of krill

The thelycum of females has been described in the species listed in Table I1 (p. 379). Guglielmo and Costanzo (1978) have now published further descriptions of the thelyca of species in the genus Euphausia: E . americana, E. eximia, E . mutica, E . diomedeae, E . recurva, E. superba, E . vallentini, E . lucens, E. frigida, E. pacijica, E. crystallorophias, E . tenera, E . similis, E. similis var. armata, E . mucronata, E. sibogae, E. distinguenda, E. lamelligera, E. gibba, E . gibboides, E. fallax, E . sanzoi, E . pseudogibba, E . paragibba, E . spinifera, E . hameni, E. longirostris and E . triacantha. The only species of this genus whose thelycum has not so far been described is E . nana. An examination of these descriptions of Guglielmo and Costanzo confirms that the form of the thelycum is diagnostic of the species. Its routine use for specific identification of females is still impeded by the difficulty in producing recognizable drawings of this complex structure. Further, the significance of its form relative to that of the petasma of the males, which is also diagnostic of the species, still remains unknown. Genetic variability in some euphausiids is briefly discussed on (p. 382). Ayala and Valentine (1978, 1979) and Valentine and Ayala (1 978) discuss further .the genetic variability they found within Euphausia superba, E. mucronata and E. distinguenda.

ADDENDUM

597

3. Distribution and synonymy

The following are additional references on the occurrence and distribution of species. Genus Bentheuphawia B. amblyops: Cox and Wiebe, 1979. Genus Thysanqodu T . monacantha: Ponomareva and Drobysheva, 1978 ; Cox and Wiebe, 1979.

T . tricuspidata: Cox and Wiebe, 1979. T . aequalis: Cox and Wiebe, 1979. T . orientalis: Ponomareva and Drobysheva, 1978; Cox and Wiebe, 1979.

T . acutifrons: Cox and Wiebe, 1979. Genus Meganyctiplzanes M . norvegica: Brodie et a,?., 1978; Jukic, 1978; Bliznichenko et al., 1979; Brunel, 1979; Cox and Wiebe, 1979; Pearcy et al., 1979. Genus Nyctiphanes N . cowhi: Jukic, 1978. N . australis: Fenwick, 1978; Taw and Ritz, 1979. N . capensis: Nepgen, 1979. Fenwick (1978) identifies this species off New Zealand but this requires confirmation. Genus Euphausia E. americana: Cox and Wiebe, 1979. E . lcrohni: Jukic, 1978; Blackburn, 1979; Brockmann, 1979; Cox and Wiebe, 1979. E. brevis: Cox and Wiebe, 1979. E . superba: Dzik and Jazdzewski, 1978; Jazdzewski et al., 1978; Kalinowski, 1978; Naumov, 1978; Fevolden, 1979; Fiacher, 1979; Hempel et al., 1979a, b; Nast, 1979; Takahashi, 1979; Warner, 1979.

E. vallentini: Ponomareva and Drobysheva, 1978. E . lucens: Ponomareva and Drobysheva, 1978; Nepgen, 1979. E . frigida: Dzik and Jazdzewski, 1978; Ponomareva and Drobysheva, 1978; Fevolden, 1979. E . Pacijica: Odate, 1978; Pieper, 1979; Pogodin, 1979. E . crystallorophim: Ponomareva and Drobysheva, 1978; Fevolden, 1979; Hempel et at., 1979a.

698

THE BIOLOQY OF EUPFIAUSIIDS

E. tenera: Cox and Wiebe, 1979. E. gibboides: Pugh, 1972; Cox and Wiebe, 1979. E. paragibba: the records of Ponomareva and Drobysheva (1978) are very far south and require confirmation. E. hemigibba: Jukic, 1978; Odata, 1978; Cox and Wiebe, 1979. E. triacantha: Dzik and Jazdzewski, 1978; Fevolden, 1979. Genus ThysanoZssa T. longipes: Pogodin, 1979. T.inermis: Zelicman et al., 1978, 1979; Bliznichenko et a,?., 1979; Brunel, 1979; Cox and Wiebe, 1979; Hopkins and Evans, 1979; Pearcy et al., 1979; Pogodin, 1979. T. longicaudata: Bliznichenko et al., 1979; Cox and Wiebe, 1979. T. parva: Cox and Wiebe, 1979. T. gregaria: Cox and Wiebe, 1979. T.vicina: Dzik and Jazdzewski, 1978. T.nzacrura: Dzik and Jazdzewski, 1978; Ponomareva and Drobysheva, 1978; Fevolden, 1979; Makarov, 1979a. T. rmchi: Zelicman et al., 1978, 1979; Bliznichenko et al., 1979; Brunel, 1979; Hesthagen and Gjermundsen, 1979; Hopkins and Evans, 1979; McConnaughey and McRoy, 1979; Pearcy et al., 1979; Pogodin, 1979. Genus Nematoscelis N. dificilis: Odate, 1978. Ponomareva and Drobysheva (1978) have reported this species in the southern hemisphere but this ie presumably an error in their paper. N. megabps: Cox and Wiebe, 1979. N. tenella: Cox and Wiebe, 1979. N. microps:Ponomareva and Drobysheva, 1978 ;Coxand Weibe, 1979. N. atlantica: Cox and Wiebe, 1979. Genus Nematobrachion N. sexspinosum: Cox and Wiebe, 1979. N. boopis: Ponomareva and Drobysheva, 1978; Cox and Wiebe, 1979.

Genus Styloeheiron S. carinaturn: Ponomareva and Drobysheva, 1978; Cox and Wiebe, 1979.

S.afine: Cox and Wiebe, 1979. 8.suhmi; Cox and Wiebe, 1979.

599

ADDENDUM

8.elongatum: Cox and Wiebe, 1979. S. longicorne: Cox and Wiebe, 1979. S. abbreviatum: Cox and Wiebe, 1979. 4.

The lakme and reproduction

The stages of development of the eggs of euphausiids as described by G. 0. Sars are illustrated in Mauchline and Fisher (1969). Free eggs of Euphaasia superba have been examined by Hempel et al. (1979a) who grouped them into four developmental stages : 2 to about 16 cells. Stage I Stage I1 many cells. form of embryo visible as dark zones within the egg. Stage I11 Stage IV limb buds visible. The time elapsing between the egg being laid and attainment of each of these stages is not yet known. Abnormal eggs occurred in the samples and, according to Hempel et al., were probably not fertilized. The nauplius and metanauplius of several Antarctic species have been examined by Makarov (1979b)and his measurements of them are given in Addendum Table I. TABLEI. ADDITIONAL DATAON BODYSIZES(mm) OF EARLYLARVAL STAGER (see TABLE111, p. 423),from Makarov (1979b) ~~

Species

Naupliua

Metanaupliua

Euphausicc superba E . fr+qida E . ergstallmophias E . tkcantha Thysanoiiaaa m m r a

0.73-0.80 0.49-0.65 0.60-0.67

0.70-0.7 8 0.46-0.62 0.67-0.67 0.61-0.73 0.46-0.56

0*42-0.60

The occurrence of the eggs of Euphausia superba bathymetrically and geographically and the breeding ecology of this species are still incompletely known. Worner (1979) found gravid females in the southern Scotian Arch and Scotia Sea during the period December to March, 1977-1978, and major concentrations of eggs in February on the shelf in the approaches to the Antarctic Sound and in the northwestern corner of the Weddell Sea. Eggs were also most common, the previous year, in the Antarctic Sound and Bransfield Strait in February, 1977 (Hempel et al., 1979a). Their numbers then decreased while those of nauplii and especially calyptopes increased to maxima in March.

600

THE BIOLOGY OF EUPHAUSIIDS

Pogodin (1979) examined samples of euphausiid eggs from the Tatarsky Strait. Thysanoessa raschi spawned from April to August, the eggs having a diameter of 0.35-0.45mm. Eggs of diameter, 0.5000.925 mm, abundant between April and June, were those of T . inerrnis while eggs of 0.425-0-625 mm, which occurred from July to October, were probably those of Euphuusia pacijku. Spawning of euphausiids in the Tatarsky Strait (50"N) starts later and the spawning period is much longer in duration than in the warmer waters of the Sea of Japan. The spawning seasons of the species in the Tatarsky Strait did not coincide. Similarly, spawning of Antarctic species commences at different times. Makarov (1979c) found that five species commenced spawning in the following succession : E . triacantha was first, followed by E . frigida, Thysanoessa macrura, Euphuusia crystallorophias and finally E . superba. This order of succession is primarily dependent upon the latitudinal region occupied by each species; northern species, and presumably also populations, spawn before southern ones. ThysanoL;ssa species developed from the egg to the furcilia stages in 31-50 days in an aquarium but Pogodin (1979) does not state the environmental temperature of the experiments. The vertical distribution of the larvae of ThysanoGsa macrura in the Scotia Sea has been described by Makarov (1979a). Nauplii and metanauplii occurred deepest, at depths of 200-1000 m while more advanced stages had their centres of distribution closer t o the surface. Thus a developmental ascent of larvae, an ontogenetic migration, takes place in this species and is similar to that described for Euphausia superba by James Marr and illustrated in Mauchline and Fisher (1969). Hempel et al. (1979a) obtained conclusive confirmation of this developmental ascent in E . superba larvae. 5. Vertical distribution and migration

Methods of catching and sampling euphausiids are discussed on pp. 436 and 550. Mohr (1979b) describes a commercial midwater trawl and the incorporation of a pump to haul Euphausia superba inboard. Matuda et al. (1978) describe a new midwater trawl while side trawling using a net and a pump to effect continuous sampling of surface concentrations of E . superba is described by Kanda et al. (1978) and Matuda et al. (1979). The influence of ambient light intensities in controlling the diurnal vertical migrations of euphausiids is discussed on p. 439. Euphuusia superba performs a diurnal vertical migration (Arimoto et al., 1979) and Takahashi (1979) found that the relation between surface light

ADDENDUM

601

intensity ( I in lux) and krill density (d in g caught/h) at the surface is: log d = 4.2 - 1.4 log I . The ambient light levels, in which sub-surface aggregations of this species occurred, appear to vary quite widely, according to Takahashi. Avoidance of direct light by euphausiids in the laboratory is well known and Arimoto and Inoue (1978) demonstrated this behaviour in E.pacifica. The use of echo sounders to study the distribution of euphausiids is briefly discussed on p. 439. Pieper (1979) determined the distribution of the biomass of E . pacijica in an area off southern California through use of a 102 kHz sounder and reviews previous feasibility studies. He determined the following linear relationships : SV= 0.89 D W - 77.83

Ti3 = 2.93 L - 106.27 TS=O.89 DBW-77.83

where DBW is 10 log mean dry weight (mg) per euphausiid in the net sample; DW is 10 log dry weight (rng dry weightlm3) of the euphausiid biomass; L is 10 log mean body length (mm) in the net sample of euphausiids; Sv is the average volume scattering strength (dB/mS); Ti3 is target strength (dB). Jukic (1978) ascribed the source of an epibenthic scattering layer detected with a 30 kHz sounder in the Adriatic Sea to populations of Meganyctiphunes norvegica, Nyctiphanes couchi and the mysid Lophogaster typicus. The association of Thysanobsa raschi and T . inerrnis with sound scattering layers in the Balsfjord, northern Norway, is discussed by Hopkins and Evans (1979) and Pearcy et ab. (1979). Cram (1978), Fischer (1979) and Mohr (1979a) surveyed concentrations of Euphuusia superba using echo sounders in the western Atlantic sector of the Antarctic. Studies of the vertical migrations of euphausiids in sonic scattering layers are studies of the behaviour of aggregations rather than of dispersed populations. Some recent studies of diurnal migration examine the degree of aggregation or dispersion present within the populations. Zelicman et ab. (1979) confirm that Thysanobsa raschi and T.inemis in the Barents Sea perform die1 vertical migrations at all seasons of the year, especially the individuals that occur in aggregations. Individuals that are outside aggregations may not be such rhythmic and active vertical migrators. Kalinowski (1978) describes

602

THE BIOLOGY OF EUPHAUSIIDS

vertical migration of Euphausia superba associated with d i w d aggregation near the surface and nocturnal dispersion a t 150 m depth. 6. Food and feeding

Kato et al. (1979) conclude from laboratory determinations of the filtering rates of Euphausia superba that they must require food other than phytoplankton to satisfy their carbon requirement. 7. Chemical composition

Bykov et al. (1978) examine the proximate composition of Euphausia superba seasonally and relative to body size and 'sex and can account for the variations in results of previous investigations. The carbon ; nitrogen ratio was found by McConnaughey and McRoy (1979) to be 4-0 in Thysanokksa raschi, the same as that already quoted in Table VIII (p. 462). The chitin and chitosan of Euphausia superba has been studied by Anderson et al. (1978). The chitin was coloured by small quantities of carotenoid pigments provisionally identified as astaxanthin, astaoin, a series of fatty acid esters of astaxanthin and a protein-bound carotenoid that was not idenbified. A further study of the structure of chitin of E . superba has been made by Galat and Popowicz (1978). Concentrations of various metal pollutants are quantified in Meganyctiphanes norvegica from the Straits of Messina by Calapaj et al. (1978). The presence of a wide range of metals in Euphausia superba and E. pacijica was detected qualitatively by Kikuchi (1979). Some were determined quantitatively in a sample of E . superba collected on 31 December, 1977 a t 63'33'5, 132"53*5'Eand in a sample of E. pacijica collected in August, 1978 in Kagoshima Bay, Japan. They contained the following concentrations, in mg/lOO g wet weight, of the following metals.

Zinc : E. superba, 0.45. Arsenic : E . superba, 0-05. Selenium: E . superba, 0.015. Cadmium: E. superba, 0.001. Mercury: total mercury in E . superba was 0.001 and in E . pacifica 0.003. Methyl mercury concentrations were 0.0005 and 0.002 respectively. Lead: E . superba, 0.01. Fluoride : fluoride has been found at high concentrations in Meganyctiphanes norvegica and Euphausia superba by Soevik and Braekkan

ADDENDUM

603

(1979). The range is 133-240mg fluoride/lOOg fat free dry weight euphausiids. This is a factor of 10-110 higher than comparable concentrations in the copepod Calanus finmarchicus Gunn. and the decapod Pandalus borealis Krcayer. The fluoride was located predominantly in the exoskeleton of Euphausia superba, the muscle tissue having concentrations of 57-75 mg fluoride/lOO g fat free dry weight. The presence of these high concentrations has been confirmed in two other laboratories, according to the authors. The fluoride in the bodies is readily available nutritionally and its concentrations are 7 times higher in freeze-dried or extracted meat and 24 times higher in the whole euphausiids than the permissible limits of 10 mg fluoride/lOO g, calculated as sodium fluoride set by the Food and Drug Administration of the United States. Consequently, krill, peeled or otherwise, does not comply with these requirements for human consumption. D D T : Concentrations of DDT residues have been measured in E . superba by Lukowski (1978) in samples from the western Atlantic sector of the Antarctic. Harding and Vass (1979), in studying uptake of p, p'-DDT directly from the water by Thysanoessa rmchi, conclude that a considerable amount must be adsorbed on particles and not metabolically available to the euphausiids. The relationship of calorific equivalent of body weight (We in cal/individual)relative to body length ( Lin mm) of tropical euphausiids, the species of which were not identifled, was determined by Shushkina and Sokolova (1972) as :

We= 0 027 L232 The specific heat and heat conductivity of fresh gravid female Euphausia superba measured immediately the animals were caught had values of 0.88 (kcal/kg "C) and 0.64 (kcal/m.h. "C) respectively (Suzuki et at., 1979). Lipids : The crude fat content of samples of Euphausia superba increased continuously throughout the season November to February, being as low as 11% in November and as high as 22-28% dry weight in February (Yanagimoto et ab., 1979a). Water content ranged from 73% t o 79% wet weight; crude protein from 60% to 73% dry weight and ash from 11% to 15% dry weight. Ciliatine, 2-aminoethylphosphonic acid, has been isolated from the lipids of E. superba by Tamari (1979). Reference to the occurrence of proteases responsible for autolysis of fresh krill is made on p. 477. Nagayama et al. (1979) has studied the action of a further three enzymes involved in the deterioration of hrill ; these are lipase, carboxylesterase and catecholoxidase. The combined

604

TRE BIOLOQY OF EUPHAUSIIDS

effect of temperature and low pH inactivated krill protease, so reducing rates of autolysis (Kubota and Sakai, 1978). 8. Vision and bioluminescence

Image formation in euphausiid eyes is effected by a refracting superposition mechanism as shown by Land et al. (1979). This mechanism is different from that found in caridean decapods where image formation is effected not by refracting cones but by mirrors (Land, 1978). The structure of the eye of Thysanopoda tricwpidata is described by Meyer-Rochow and Walsh (1979)while those of Meganyctiphanes norvegica, Nyctiphunes simplex, Euphausia paci$ca, Nematoscelis megalops, N . microps, N . atlantica, Nematobrachion boopis, Stylocheiron longicorne and S. maximum are described by Land et al. (1979). Land and Burton (1979)conclude from all these studies that the eyes of euphausiids have an optical system closely similar to that of nocturnal insects. Portable equipment for use on board ship to scan the bioluminescence of marine organisms spectrally is described by Collier et al. (1979). Emission spectra of homogenized photophores of Thysanopoda monacantha and Meganyctiphanes norvegica are presented graphically. Cram and Malan (1977) discuss the possibility of locating surface and sub-surface swarms of euphausiids by remote sensing techniques utilizing the spectral emission bands of their bioluminescence. 9. Internal anatomy and physiology

The body density of Meganyctiphunes norvegica and Euphausia superba has been found by Kils (1979a) to increase with increasing body length of the animal. It is 1.042 g/ml for a 10 mm euphausiid and 1.068 g/ml for a 60 mm euphausiid. The relationship is : = 1.028 (1

- 3.38 x

L0*590)-1

where Q is density and L is body length. Greenlaw (1977) found that the density of E . pacijica is 1.037-1-052g/ml. Suzuki et al. (1979) estimated the density of E . superba to be 1.01-1.04 g/ml. The sinking speed of E . superba and Meganyctiphunes norvegica is described by the equation : 8 = 0-0701 L1*07 where S is the sinking speed in cm/s and L is body length in mm (Kils, 1979b). Larger animals have greater sinking speeds than smaller

ADDENDUM

605

animals. Consequently, t,hey have t o expend more energy tq maintain their bathymetric position in the water column. A 60 m m Euphausia superba has to expend 61 times more energy per unit body weight than a 6 mm individual in order to prevent itself from sinking. The equation describing the amount of energy required by euphausiids of different body length to maintain their vertical position is :

P/W , = 1.76 L1.66 where P/W , is energy per unit wet weight in joules/day/mg wet weight and L is body length in mm (Kils, 1979b). Kils (1979~)examines the mechanism of swimming in Meganyctiphanes norvegica. The rhythm of pleopod beating is described along with swimming speeds and behavioural patterns relating to swimming. Speeds of up to 8 times the body length of the animal per second can be achieved by the normal process of forward locomotion induced by metachronal beating of the pleopods. The “escape reaction”, effected by flexion of the abdomen inducing backward motion, can result in speeds of up t o 11 times the body length of the animal per second. Shushkina and Pavlova (1973) examined the metabolic rate of tropical euphausiids, species not named, relative to other phylogenetic groups of planktonic organisms. They found that

T = 0.416 Pg1’ where T is the respiration rate expressed as mcal/day/individual and M is body weight expressed as mcal/individual. A similar equation

T = 0.066 Jf0.052 was found by Musaeva (1978) for south-western Pacific species that were not identified. Lipskaya and Drobysheva (1978, 1979) measured the respiration rates of Euphausia lucens, E . recurva, E . paragibba, E. crystallmophias and E . triacantha at 20°C and fitted %hefollowing equation : R=O-26 W@*l

where R is pl O,/animal/h and W is the body wet weight in mg. Segawa et al. (1979) found that E . superba consumed 0.049-0.188 pl 02/mgwet weight/h a t - 1.5-5°C. Further measurements of respiration rates of this species are given by Rakusa-Suszczewki and Opalinski (1978). The possibility that the mitochondria1 and microsomal electron transport system (ETS) may provide an indirect method of measuring respiration is discussed briefly on p. 488. Bamstedt (1979) examined the relationship between ETS and respiratory rate on the one hand

606

THE BIOLOGY OF EUPHAUSTIDS

and body weight on the other in a selection of planktonic organisms including Meganyctiphunes norvegica and Thysanoha species. Both were related to body weight and both varied considerably with season, maxima usually occurring in the spring and minima in the summer. The ETS activity, however, was more variable than the respiratory rate and seasonal variations in the two were poorly correlated. Consequently, ETS activity may not be satisfactory for predicting population metabolism. Euphausia superba requires high oxygen saturation levels in its environment, Kils (1979b) found an LD,, for this species between 80% and 85% oxygen saturation. The tolerance of E . pacijica to thermal shocks of short duration, such as would be encountered in passage through the condenser cooling systems of thermal nuclear power plants, has been examined by Craddock (1979). Environmental temperatures in which E . paciifica lives in Puget Sound usually range from 7°C to 11°C. Exposure to temperatures of 23"-24"C for 15 minutes killed 1 1 4 3 % of the animals tested. 10. Growth, maturity and mortality

Various biometrical relationships additional t o those in Table XV (p. 491) are now available and are shown in Addendum Table 11. Brood size increases with increasing body size within a species of euphausiid (p. 498). Jazdzewski et al. (1978) found that the number of eggs in the ovary of Euphausia superba varies from 2300 to 13 500. The following equations describe the number of eggs ( N ) in the ripe ovary relative to body length ( L in mm) and body volume (V in pl).

N = 0.0001

L44'Ss

N = 0.3108

J'1.s455

The weight and volume of the ripe ovary in this species represents 30-50% of the females weight and volume. The ripe ovary of E . superba is consequently much larger relative to the body than in other euphausiids where it represents about 10% of the body volume (p. 495). The following equation relates the volume of the ripe ovary, V o in pl, t o the femde"s body length, L in mm -.

VTo = 0.0003 L3*sz0s Growth of E . superba has been studied in the laboratory by Murano et al. (1979). The regression equation relating the duration of the

607

ADDENDUM

intermoult period (IPin days) to total body length ( L in mm) is : log IP= 1.157+0.017 L Growth factors observed in the laboratory were, on average, small and reflect deficiencies in the food and conditions of maintenance provided rather than the true growth factors in the environment. Continuous breeding throughout the year in some species of euphausiids is discussed on p. 512. Taw and Ritz (1979) state that TABLE11. REQRESSION CONSTANTS OF BIOMETRICAI. RELATIONSHIPS (Jazdzewski et al., 1978; Kils, 1979a, b ) Regression comtants y=bx+a or log y = b log x + a Species E u p h a k a superba Juveniles and males gravid females all individuals juveniles males females gravid females all individuals all individuals Meganyctiphanes norvegica

Parameters

log V on log TL log W, on log T L

log W, on log T L Euphawia superba log W,, Meganyctiphanes norvegica

on log

TL

a

b

-2.17 - 2.49 - 2.44 - 2.52 - 2.16 - 2.96 - 2.47 - 2.74 - 2-80 - 2.17 - 2.85 - 4.16 - 4.00 - 4.76 - 4.99

3.00 3-29 3.16 3.23 3.03 3.52 3-27 3.38 3.40 3.00 2-98 3.76 3.80 3.67 3.82

T L , total length, base of eye to end of telson; V , body volume; W,, body dry weight; Ww,body wet weight; W,,, underwater body wet weight.

calyptopes and furciliae of Nyctiphanes australis are found throughout the year off south-eastern Tasmania, so indicating continuous breeding within these populations. Saprykina (1979) examines the cycles of gonad development and degeneration in Euphawia pacifica, Thysano&sa raschi and T . inermis from the northern Sea of Japan. The cycle can be divided into generative and resting periods. The generative period includes the following stages : ( l ) ,formation of secondary sexual characters; (2), maturation of the gonads in males, final formation of

608

THE BIOLOGY OF EUPHAUSIIDS

external sexual characters, growth of the ovary in females; (3), mating, final maturation of the females; (4), spawning of the females. The resting period includes : (5), final spawning; (6), post-spawning restoration of the gonads; (7), recurrent maturation in males and re-growth of the ovary in females; (a), mating, final re-maturation of the ovary; (9), spawning. Euphausia paci$ca breeds seasonally in the northern Sea of Japan, first spawning occurring a t a body length of 16-18 mm. A proportion of the population survives to breed a second time a t an age of 2 years (Saprykina, 1979). This cycle is very different from that described in this species in Table XXI (p. 51 1). Length frequency distributions in samples of Euphausia superba, E. crystallorophias, E . frigida and Thysanoessa rnacrura in the western Atlantic Sector of the Antarctic are given by Fevolden (1979). The data are inconclusive as regards age and size a t sexual maturity. Jazdzewski et al. (1978) also provide length-frequency histograms of Euphausia superba in various regions of this sector. Considerable variation is apparent and few conclusions on age structure can be drawn. 11. Ecology of distribution

Arctic Ocean and Peripheral Seas. Further information on the distribution of Thysanohsa raschi, T . inermis and Meganyctiphanes norvegica in the Norwegian and Barents Seas in 1976-1977 is given by Bliznichenko et al. (1979). North Atlantic, north of about 40"N. Estimations of environmental population densities of euphausiids off Nova Scotia made by Brodie et al. (1978) take account of results of net sampling, sonar surveys and the contents of whale stomachs. The Baltic Sea. Thysanoessa raschi is recorded in the Kiel Bight by Hesthagen and Gjermundsen (1979). Atlantic Ocean, 4O"N-4O0S. The occurrence and origin of the euphausiid fauna of the Middle Atlantic Bight between about 42"N and 35"N off the American coast has been examined by Cox and Wiebe (1979). Some 31 species were found and include Arctic-boreal species from shelf waters north-east of Cape Cod, transition zone species from the adjacent slope water and tropical-subtropical species originating from the Gulf Stream and Sargasso Sea. They discuss the structure of the region, mechanisms of dispersion and movement and the overwintering of warm-winter species in this rigorous area. PaciJic Ocean, off Japan. The euphausiid fauna of the Kuroshio Current region is described seasonally by Kuznetsova (1977) but this

ADDENDUM

609

paper has not been examined as yet by the author. Odate (1978) describes the distribution of the species off north-western Japan. Indian Ocean. The distributional data of Baker (1965), quoted by Mauchline and Fisher (1969), on Euphausia species in the Indian Ocean have been re-analysed by Williamson (1978) to demonstrate a new method of data processing in principal components analyses. Antarctic Ocean and South Australia. The distribution of species in the region of southern and eastern Australia and extending southwards to the ice edge in the Antarctic is described by Ponomareva and Drobysheva (1978). They identify Nemutoscelis dificilis in latitudes 20"-33°S and Euphausia paragibba as far south as 45%. Nematoscelis dificilis is restricted to the North Pacific and so this new record is probably in error. Euphausia paragibba has not previously been recorded south of about 20"s and consequently its occurrence as far south as 46;"s is also questionable. The reproductive ecology and distribution of the larvae of Thysanoessa macrura in the Scotia Sea has been investigated and discussed by Makarov (1979a). Euphausia superba and E. frigida appear to have breeding populations in the region of Bouvet Island (Fevolden, 1979). 12. Predators and parasites

Predation of euphausiids by fish is reviewed on p. 529. The snake blenny, Lumpenus lampretaeformis (Walbaum), occasionally eats euphausiids in western Scotland (Gordon and Duncan, 1979). Cod, capelin and herring in the Balsfjord, northern Norway, feed on Meganyctiphanes norvegica, Thysan&ssa raschi and T . inermis to a large degree (Pearcy et aE., 1979). Another inshore fish, the American sand lance, Ammodytes americanus De Kay, feeds on euphausiids to a small extent off Cape Cod, Massachusetts (Meyer et al., 1979) as does the snoek, Thyrsites atun (Euphrasen), on Euphausia lucens and Nyctiphanes capensis off South Africa (Nepgen, 1979). One unidentified euphausiid occurred among the stomach contents of 28 Atlantic salmon, Salmo salar L., caught at Port Burwell, Northwest Territories, Canada by Neilson and Gillis (1979). In the Mediterranean, the hake, Merluccius merluccius L., and the horse mackerel, Trachurus trachurus L., were found by Jukic (1978) to be feeding almost exclusively on Meganyctiphanes norvegica and the mysid Lophgaster typicus during daylight hours. A few Euphausia krohni and E. hemigibba were also present among the stomach contents. In the Bering Sea, juvenile walleye pollack, Theragra chalcogramma (Pallas), feed principally on copepods and unidentsed euphausiids

610

THE BIOLOGY OF EUPHAUSIIDS

while the adults feed principally on euphausiids, amphipods and fish (Bailey and Dunn, 1979). Porichthys notatus Girard, Cithurichthys sordidus (Girard), Anaplopoma Jimbria (Pallas), Merluccius productus (Apes), and Prionace glauca (L.) prey on Euphausia paci$cu and Thysanoessa spinifera in Monterey Bay, California (Morejohn et al., 1979). The Peruvian jack mackerel Trachurus symmetricus murphyi (Nichols) includes euphausiids as a major portion of its diet (Konchina, 1978). The ribbon fishes Trichiurus japonicus and T . lepturus L. feed occasionally on euphausiids off Taiwan (Lee, 1978). The importance of micronektonic fish as predators of euphausiids is discussed on p. 530. The following additional information is available. Stenobrachius leucopsarus (Eigenmann and Eigenmann) feeds t o a large extent on Euphuusia paciJica off California (Collird, 1970 ; Cailliet, 1972). Gorelova (1974, 1978) did not identify the species of euphausiids that were eaten by Myctophum spinosum (Steindachner), M . nitidulum Garman, M . brachygnathos (Bleeker), M . aurolaternutum Garman, Ceratoscopelus warmingi (Lutken) and Bolinichthys longipes (Brauer). Other myctophids, found by Collard (1970), feed on Euphuusia paciJica, E . hemigibba and Nematoscelis di$cilis in the eastern Pacific and are: Diaphus lucens, D . theta Eigenmann and Eigenmann, Diogenichthys laternatus (Garman), Lampanyctus australis, L. macropterus (Brauer), L. regalis (Gilbert), L. ritteri Gilbert, Myctophum spinosum, Parvilux ingens, Symbolophorus californiensis, Tarletonbeania crenularis (Jordan and Gilbert) and Triphoturus mexicunus (Gilbert). The Antarctic species Electrona antarctia (Giinther) feeds predominantly on Euphausia superba; a detailed study of the selection of adults and larvae by different sized Electrona antarctica is described by Rowedder (1979).

Macrourid fish are also predators of euphausiids as described on p. 532. Macrourus berglax Lacdpede consumes some euphasiids (Geistdoerfer, 1979) while Coryphaenoides rupestris Gunn. eats many (Podrazhanskaya, 1968; Buit, 1978) in the north Atlantic. Buit also found that Glyptocephalus cynoglossus (L.) and Coelorhynchus coelorhynchus (Risso) eat euphausiids to some degree. Macpherson (1979) found that Hymenocephulus italicus Giglioli, Nezumia aequulis (Gunther) Coelorhynchus coelorhynchus and Trachyrhynchus trachyrhynchus (Risso) feed to some extent on euphausiids, principally on Meganyctiphanes norvegica, in the western Mediterranean. Marshall and Merrett (1977) found euphausiids among the food of Hymenocephalus italicus and Chbrophthalmus agassizi Bonaparte off north-west Africa. The importance of euphausiids in the diets of macrourids is emphasized by their occurrence in the diets of 24 of the 25 species examined by Okamura

ADDENDUM

61 1

(1970). Haedrich and Polloni (1976)foundthat Coryphaenoides carapinus Goode and Bean includes a few euphausiids in its diet. Predation of Antarctic krill by fish is described on p. 533. Notothenia rossi marmorata (Fischer) feed exclusively on Euphuusia superba while Champsocephulus gunnari Lonnberg off South Georgia, where Euphausia superba was scarce, substituted amphipods and to a smaller extent mysids (Kock, 1979). One further fish, Pseudochuenichthys georgianus Norman fed on Euphuusia superba but also, to an equal extent, on fish. Feeding of other Antarctic fish on E . superba is noted by Rembiszewski et al. (1978). Micromesistius australis Norman, the southern blue whiting, feeds predominantly on Euphausia species in the south-western Atlantic (Otero, 1977). The short-finned squid Illex illecebrosus (Le Sueur) consumes Meganyctiphunes norvegica in the Bay of Fundy (Brown et al., 1979) while the market squid, Loligo opalescens Berry, feeds primarily on Euphuusia pmifica and Thysanoessa spinifera in Monterey Bay, California (Karpov and Cailliet, 1979). The following birds in Monterey Bay, California feed on Euphausia pacijica and Thysanoessa spinifera according to Morejohn et al. (1979): Pufinus griseus (Gmelin), Larus heermanni and Uria aalge. New Zealand birds feed on surface swarms of Nyctiphanes species (Fenwick, 1978) : Pufinus griseus, Daption capensis L., Diomedea bulleri Rothschild and Larus novaehollandiae. The Antarctic burrowing petrels, Pachyptila desolata (Gmelin) and Halobaena caerulea Gmelin, feed principally on Euphausia superba but also include, a range of other crustaceans as lesser components of the diet, including Thysanoessa macrura (Prince, 1980). Birds are dependent upon the occurrence of surface’swarms of euphausiids in order to gain access to them. The greater shearwaters, Pufinus gravis (O’Reilly), feed on surface swarming Meganyctiphanes norvegica in the Bay of Fundy (Brown et al., 1979). 13. Euphausiids in the marine economy

The potential exploitation commercially of Euphuusia superba has resulted in the development of the relevant technology (Bykov, 1978; Tsareva et al., 1978; Kanda and Hotani, 1979). Information on midwater trawls, echo sounding and other aspects of location and capture of E . superba are given by Cram and Malan (1977), Bogdanov and Lyutimova (1978), Cram (1978), Lestev (1978) and Hempel et aE. (1979). Yanagimoto et al. (1979b) describe a method of eviscerating krill by application of water jets. Toyama and Yano (1979) use a juice extractor to dispense with the exoskeleton prior to processing. Methods

612

THE BIOLOGY OF EUPHAUSIIDS

of processing and development of products of krill are described by Gamygin et al. (1978), Govorunova and Gun'ko (1978), Roschke et al. (1978) and Schreiber et aE. (1979). The technology of canning krill and the quality of the canned products has been investigated by Tanaka et al. (1978, 1979a). Vitamin content of processed krill is estimated by Secomska and Jelinska (1979). Freezing and defrosting of krill has been studied by Tanaka et al. (1979b) and heat transfer coefficients determined. Separation and concentration of protein fractions from fresh and defrosted krill are described by Toyama et al. (1979). Insoluble protein fractions are recovered during the production of soluble fractions by methods given by Watanabe et al. (1979). The nutritional value of these proteins is defined by Kunachowicz et al. (1978). The thermophysical characteristics of krill juice are described by Gromov and Koroleva (1978). The total biomass of Euphausia superba and its annual production have been re-estimated by Tom0 and Marshoff (1977)as 220-440 million metric tons and 130-660 million tons per year respectively. Hirayama et al. (1979), however, estimate an even greater biomass-360-1370 inillion tons. The realistic estimation of these parameters of the stocks of E . superba is one of the primary aims of the multi-national programmes of Biological Investigations of Marine Antarctic Stocks and Systems (BIOMASS, 1977, 1978). The first experiments of these investigations (FIBEX) are scheduled for February 1981. They are primarily feasibility studies to examine the use of sonar techniques to estimate and map quantitatively the distribution of E . superba in the western Atlantic sector. The investigations are planned t o extend over a number of years and to examine the marine Antarctic ecosystem as a whole. The euphausiids, especially E . superba, form a major co&ponent of this ecosystem and so much new knowledge on these euphausiids should be obtained. A commercial fishery for euphausiids must of necessity exploit aggregations (pp. 524, 550). The Japanese commercial fishery on the Sanriku and Joban coasts of northeastern Honshu for E . paci$ca, locally known as tsunonashi-okiami, is described by Odate (1979). This fishery commenced in the 1950s. Landings have increased from 8662 tons in 1971 to 41 215 tons in 1978. The aggregations of E . paci$ca are associated with extensions of cold water, less than 5°C at 100 m depth, in the area between 35"N and 42"N, west of 145'E during the seasonal period February to May. The increasing catches in recent years are associated with southward expansions of cold Oyashio Current waters. Aggregations associate with interphases between cold and coastal water masses.

ADDENDUM

613

The surface swarms of Meganyctiphnes norvegica that occur seasonally in July to September in the Bay of Fundy have been studied by Brown et al. (1979). Their presence appears to be associated with strong tidal streams and were most common in spring tide periods. Brown et al. suggest that vertical turbulence transports the animals to the surface and is responsible in initiating swarm formation. Small swarms of M . norvegica, similar to the spherical swarms of Nyctiphanes species described by Fenwick (1978) off New Zealand, also occur in the Bay of Fundy, but parallel orientation was present within the swarms in which densities of the order of 1000 animals/m3were estimated. Zelicman et al. (1978, 1979) studying Barents Sea species found that they occurred aggregated but that a portion of the populations lived outside aggregations. The individuals outside aggregations may, according to these authors, be less active diurnal vertical migrators that become separated from rhythmically migrating aggregations ; this may happen for purposes of feeding or moulting. Kalinowski (1 978) describes vertical migration associated with diurnal aggregation of Euphusia superba at 50 m depth and nocturnal dispersion at 150 m depth. The occurrence of E . superba in aggregations is discussed by Mohr ( 1 9 7 9 ~ ) . Rakusa-Suszczewski (1978) measured temperature, salinity, oxygen content, phosphate, nitrites and nitrate in the region of swarms but found no effect of these parameters, within the ranges that occurred, on swarming. Methods of estimating krill density and the total weight of E . superba in a surface aggregation are discussed by Matuda et al. (1979). A novel method, using a radio-controlled model aeroplane incorporating a camera to photograph surface swarms, is described by Hamada et al. (1978). The types of aggregations formed by euphausiids are discussed on p. 524. Discussions of these during the planning stages of BIOMASS, especially with Mr A. de C. Baker and Professor G. Hempel, in conjunction with the recent papers of Brown et al. (1979) and Odate (1979) have produced a general concept of aggregation. This is illustrated schematically in Addendum Fig. 1. A patch is as defined on p. 524, that is it occurs on a geographical scale, its occurrence probably controlled primarily by the physicalchemical parameters of the environment. Consequently, a patch is an aggregation within a defhed environmental region. Environmental parameters me more important than behavioural reactions between individuals in maintaining the aggregation. Densities within patches are low, maximally 1-10 individuals/m3. A shoal is not correctly defined on p. 524 because that definition s t a h that it is larger than a patch. A shoal is usually a constituent

614

THE BIOLOGY OX' EUPHAUSIIDS

.

Fro. I. A concept of a euphausiid patch containing constituent shoal, swarms (W) and schools (C).

part of a patch. It can be a large aggregation up t o several km2. Densities vary in different parts of a shoal but average densities probably range from 1 to 100 individuals/m3. A swarm is often a constituent part of a shoal. They are cohesive groups of individuals without parallel orientation. Densities are in the range 1000-100 000 individuals/m3. A school is often a constituent part of a shoal. They are cohesive groups of individuals with parallel orientation. Densities are in the range 1000-100 000 individuals/m3. The present concept of aggregation of euphausiids is as follows. Aggregation is a general term for a grouping together of the animals and does not d e h e its structure or the physical-chemicalparameters of the environment or behavioural mechanisms of the animals that might be important in initiating and maintaining the grouping. A breeding aggregation is self explanatory; it is seasonal in nature and disperses after the population has bred. A patch is an aggregation effected by the physical-chemical parameters of the environment and occurs in association with the bathymetry of the seabed in coastal regions, and with fronts, undersea mounts, the ice edge and a t other environmental discontinuities in oceanic regions. Among these, of course, must be included irregularities in the distribution of phytoplankton. The average densities of euphausiids in patches are probably below threshold densities that initiate social behaviour and consequent formation

ADDENDUM

615

of shoals, swarms and schools. The combination of the micro-hydrographical features of the environment of a patch and behavioural characteristics of the euphausiids such as their association with an isolume during vertical migration may generate the threshold densities initiating formation of shoals, swarms and schools. Shoals may be formed first and a further process of microconcentration generate swarms and schools. Swarms, in which there is no parallel orientation of the animals, are immobile relative to the water whereas schools, with parallel orientation of individuals, are actively moving relative to the water. Schools can act as self-sorting mechanisms, being selective for size groups. Swarms and schools must disperse to allow the individuals to feed. Densities within shoals may be sufficiently low to be compatible with phytoplankton grazing. Dynamical studies are required on the integrity of aggregations in the contexts of time, vertical migration, feeding, breeding and moulting.

REFERENCES Anderson, C. G., Pablo, N. de and Romo, C. R. (1978). Antarctic krill ( E u p h a w i a superba) as a source of chitin and chitosan. I n “Proceedings of the First International Conference on Chitin/Chitosan” (Eds R . A. A. Muzzarelli and E. R. Pariser), pp. 54-63. Massachusetts Institute of Technology, Cambridge, USA. Arimoto, T. and Inoue, K. (1978). Study concerning phototaxis and mesh selection for Euphausia pacifica. Transactions of the Tokyo University of Fisheries, No. 2, 45-51. Arimoto, T., Matuda, K., Hamada, E. and Kanda, K. (1979). Die1 vertical migration of krill swarm in the Antarctic Ocean. Transactions of the Tokyo University of Fisheries, No. 3, 93-97. Ayala, F. J. and Valentine, J. W. (1978). Genetic variation and resource stability in marine invertebrates. I n “Marine Organisms, Genetics, Ecology, and Evolution” (Eds B. Battaglia and J. A. Beardmore), pp. 23-51. Plenum Press, New York and London. Ayala, F. J. and Valentine, J. W. (1979). Genetic variability in the pelagic environment : a paradox. Ecology, 60, 24-29. Bailey, K. and Dunn, J. (1979). Spring and summer foods of walleye pollock, Theragra chalcogramnuz, in the eastern Bering Sea. Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 77, 304-308. Bamstedt, U. (1979). Seasonal variation in the respiratory rate and ETS activity of deep-water zooplankton from the Swedish west coast. I n “Cyclic Phenomena in Marine Plants and Animals” (Eds E. Naylor and R. 0. Hartnoll), pp. 267-274. Pergamon Press Ltd, Oxford. BIOMASS (1977). Biological Investigations of Marine Antarctic Systems and Stocks. Vol. I. Research Proposals. SCAR/SCOR. Group of Specialists on Living Resources of the Southern Oceans. 1-79.

616

THE BIOLOQY OF EUPHAUSIIDS

BIOMASS (1978). Meeting of the SCAR/SCOR Group of Specialists on Living Resources of the Southern Oceans, Kiel, F.R.G., May, 1978. Biomaes Report Series, 1, 1-31. Blackburn, M. (1979). Zooplankton in an upwelling area off northwest Africa: composition, distribution and ecology. Deep-sea Research, 26A,41-56. Bliznichenko, T. E., Degtereva, A. A., Drobysheva, S. S., Nesterova, V. N. and Ryzhov, V. M. (1979). Plankton investigations in thc Baraits and Norwegian Seas in 1977. Annales Biologiques, 34 (1977), 93-94. Bogdanov, A. S. and Lyubimova, T. G. (1978). Soviet krill investigations in the Southern Ocean. Rybnogo Khozyaistva, Morskogo, No. 10, 6-9. Brockmann, C. (1979). A numerical upwelling model and its application to a biological problem. Meeresforschung, Reports on Marine Research, 27, 137-146. Brodie, P. F., Sameoto, D. D. and Sheldon, R. W. (1978). Population densities of euphausiids off Nova Scotia as indicated by net samples, whales stomach contents, and sonar. Limnology and Oceanography, 23, 1264-1267. Brown, R . G. B., Barker, S. P. and Gaskin, D. E. (1979). Daytime surface swarming of Meganyctiphanes norvegica (M. Sars) (Crustacea, Euphausiacea) off Brier Island, Bay of Fundy. Canadian Journal of Zoology, 57, 2285-2291. Brunel, P. (1979). Seasonal changes of daily vertical migrations in a supra. benthic cold-layer shelf community over mud in the Gulf of St. Lawrence. I n “Cyclic Phenomena in Marine Plants and Animals” (Eds E. Naylor and R. G. Hart,noll), pp. 383-390. Pergamon Press, Oxford. Buit, M.-H. du (1978). Alimentation de quelques poissons teleosteens de profondeur dans la zone du seuil de Wyville Thomson. Oceanologia Acta, 1, 129-134. Bykov, V. P. (1978). Main results of technological studies on krill. Rybnogo Khozyaistva, Morskogo, No. 10, 60-64. Bykov, V. P., Storozhuk, A. Ya., Radakova, T. N., Solomatina, L. F. end Sidorova, E. M. (1978). The chemical composition of krill. Rybnogo Khozyaistva, Morskogo, No. 10, 69-73. Cailliet, G. M. (1972). The study of feeding habits of two marine fishes in relation t o plankton ecology. Transactions of the American Microscopical Society, 91, 88-89. Calapaj, R., Ciraola, L., Magazzu, G. and Berdar, A. (1978). Le contenu enHg, Pb, Cd, Cr, Cu, Fe, Zn de certains poissons bathyphiles et d’un crustace du Detroit de Messine. Revue Internationale Ochnographie Medicale, 51/53, 127-137. Collard, S. B. (1970). Forage of some eastern Pacific midwater fishes. Copek, 348-354. Collier, A. F., Burnham, R . J. P. and Herring, P. J. (1979). A system for the collection of comparative emission spectra suitable for shipboard use. Journal of the Marine Biological Association. of the United Kingdm, 59, 489-495. Cox, J. and Wiebe, P. H. (1979). Origins of oceanic plankton in the Middle Atlantic Bight. Estuarine and Coastal Marine Science, 9, 509-527. Craddock, D. (1979). Effect of thermal increases of short duration on survive1of Euphausia pacifica. Fishery Bulletin, National Oceanic Atmospheric Adminktration of the United States, 76, 895-900. Cram, D. L. (1978). South African research into Antarctic krill. The South African Shipping News and Fishing Industry Review, July, 39-43.

ADDENDUM

617

Cram, D. L. and Malan, 0. G. (1977). On the possibility of surveying krill (Euphausia superba Dana) ‘in the Southern Ocean by remote sensing. South African Journal of Antarctic Research, 7 , 1-6. Dzik, J. and Jazdzewski, K. (1978). The euphausiid species of the Antarctic region. .polski5 Archiwum Hydrobiologii, 25, 589-605. El-Sayed, S. (1978). BIOMASS, a blueprint to avoid ecological tragedy in the Southern Ocean. Frontiers, 43, 38-39. El-Sayed, S. and McWhinnie, M. A. (1979). Antarctic krill: protein of the last frontier. Oceanus, 22, 13-20. Fenwick, G. D. (1978). Plankton swarms and their predators a t the Snares Islands. New Zealand Journal of Marine and Freshwater Research, 12, 223224. Fevolden, S. E. (1979). Investigations on krill (Euphausiacea) sampled during the Norwegian Antarctic Research Expedition 1976-1977. SarSia, 64, 189198. Fischer, von W. (1979). Lottechnische Aufnahme der Krillverbreitung. Archiv fiir Fischereiwissenschaft, 30, 68-70. Galat, A. and Popowicz, J. (1978). Study of the Ramen scattering spectra of chitins. Bulletin de 1’Acade&e Polonaise des Sciencm, Sdrie des Sciences Biologiqum, 26, 519-524. Gamygin, E. A., Podoskin, A. G. and Kanid’ev, A. N. (1978). Krill meat as a component of pelleted feeds for trout. Rybnogo Khozyaistva, Morskogo, No. 10, 22-24. Geistdoerfer, P. (1079). Recherches sur l’alimentation de Macrourm berglax Lacepede 1801 (Macrouridae, Gadiformes). Anndes de l’Imtitut Ockanographique, 55, 135-144. Gordon, J. D. M. and Duncan, J. A. R. (1979). Some notes on the biology of the snake blenny. Lumpenus lampretaeformis on the west coast of Scotland. Journal of the Marine Biological Association of the United Kingdom, 89, 413-41 9. Gorelova, T. A. (1974). Zooplankton from the stomachs of juvenile lantern fish of the family Myctophidae. Oceanology, 14, 575-580. Gorelova, T. A. (1978). The feeding of the lantern fishes Ceratoscopelus warmingi (Lutken) and Bolinichthys longipes (Brauer) of the family Myctophidae in the western equatorial part of the Pacific Ocean. Voprosg Ikhtiologii, 18, 673-683. Govorunova, V. V. and Gun’ko, A. F. (1978). Frozen krill as food for reared sturgeon. Rybnogo Khozyaiatva, Morskogo No. 10, 30-31. Greenlaw, C. F. (1977). Backscattering spectra of preserved zooplankton. Journal of the Acoustical Society of America, 62, 44-52. Gromov, M. A. and Koroleva, E. I. (1978). Thermophysical characteristics of krill juice. Rybnogo Khozyaistva, Morskogo, No. 10, 75-76. Guglielmo, L. and Costanzo, G. (1978). Diagnostic value of the thelycum in euphausiids. 11. Oceanic species. Genus Euphausia Dana, 1852. Archiwio di Oceanografia e Limnolog.ia, 19, 143-155. Haedrich, R. L. and Polloni, P. T. (1976). A contribution to the life history of a small rattail fish, Coryphaenoides carapinus. Bulletin of the Southern California Academy of Sciences, 7 5 , 203-211. Hamada, E., Koike, Y., Saito, K. and Suzuki, H. (1978). On a ra&o controlled inode1 airplane for observing swarms of krill. Transactions of the Tokyo University of Fisheries, No. 2, 33-44.

618

THE BIOLOGY OF EUPHAUSIIDS

Harding, G. C. H. and Vass,,W. P. (1979). Uptake from seawater and clearance of p, p’-DDT by marine planktonic Crustacea. Journal of the Fisheries Research Board of Can&, 36, 247-254. Hempel, I., Hempel, G. and Baker, A. deC. (1979a). Early life history stages of krill (Euphauaia superba) in Bransfield Strait and Weddell Sea. Meeresforschung, Reports on Marine Research, 27, 267-281. Hempel, G., Sarhage, D., Schreiber, W. and Steinberg, R . (Eds) (1979b). Antarctic Expedition 1977178 of the Federal Republic of Germany. Archiv fGr Fischereiw~ksenschaft,30, 1-1 19. Hesthagen, I. H. and Gjermundsen, B. (1979). Late summer diurnal migration in the hyperbenthos of Vejsnas Rinne, western Baltic. Meeresforschung, Reports on Marine Research, 27, 19-26. Hirayama, N., Yamada, S., Sakurai, H. and Sakuramoto, K. (1979). Stock assessment of Antarctic krill by records of a fish finder. Tramactiom of the Tokyo University of Fisheries, No. 3, 71-81. Hopkins, C. C. E. and Evans, R . A. (1979). Diurnal and horizontal variations in a zooplankton sound scattering layer. I n “Cyclic Phenomena in Marine Plants and Animals” (Ed8 E. Naylor and R. G. Hartnol), pp. 375-382. Pergamon Press, Oxford. Jazdzewski, K., Dzik, J., Porebski, J., Rakusa-Suszczewski, S., Witek, Z. and Wolnomiejski, N. (1978). Biological and populational studies on krill near South Shetland Islands, Scotia Sea and South Georgia in the summer 1976. Polskie Archiwum Hydrobiologii, 25, 607-631. Jukic, S. (1978). Contribution to the knowledge of the relationship between formation of deep scattering layer (DSL) and biological components in the central Adriatic. Acta Adriatica 19, 3-15. Kalinowski, J. (1978). Vertical migration of krill in the region of South Georgia, February-March, 1976. Polskie Archiwum Hydrobiologii, 25, 573-683. Kanda, K. and Hotani, H. (1979). Activities of the Umitaka Maru 111 research expedition for Antarctic krill fishery. Tramactions of the Tokyo University of Fisheries, No. 3, 1-14. Kanda, K., Takeuchi, S., Koike, A., Matsuda, K., Inoue, K., Akizawa, H. and Takasu, K. (1978). A preliminary experiment on the fish pump for catching live krill. Transactions of the Tokyo University of Fisheries, No. 2, 27-32. Karpov, K. A. and Cailliet, G. M. (1979). Feeding dynamics of Loligo opalescem. United States Department of Pish and Game, Fish Bulletin, 169,46-65. Kato, M.,Murano, M. and Segawa, S. (1979). Estimation of the filtering rate of the Antarctic krill under laboratory conditions. Tramactions of the Tokyo University of Fisheries, No. 3, 107-112. Kikuchi, T. (1979). Hazardous metal content in Antarctic krill. Tramactione of the Tokyo University of Fisheries, No. 3, 161-164. Kils, U. (1979a). Preliminary data on volume, density and cross section area of Antarctic krill, Euphawia superba. Results of the 2. Antarctic Expedition 1977178 of the Federal Republic of Germany. Meeresforschung, Reports on Marine Research, 27, 207-209. Kils, U. (197913). Schwimmverhalten, Schwimmleistung und Energiebilanz des antarktischen Krills, Euphawia superba. Berichte a m dem Imtitut fiir Meereskunde an der Christian-Albrechts- UniversitLit Kiel, No. 66, 1-79. Kils, U. ( 1 9 7 9 ~ ) .Swimming speed and escape capacity of Antarctic krill, Euphawia superba. Meeresforschung, Reports on MarineResearch, 27, 264266.

ADDENDUM

619

Kock, von K.-H. (1979). Fischereibiologische Untersuchungen an Fischen. Archiv f u r Fischereiwissenschft, 30, 7 1-84. Konchina, Yu. V. (1978). Some data on the nutrition of the Peruvian jack mackerel Trachurus symmetricua murphyi (Nichols). Voprosg Ikhtiologii, 18, 774-777. Koniecka, D., Wojciechowska, W. and Brzeska, Z. (1977). “Bibliography on Krill and Fish Resources in the Antarctic”, pp. 1-159. Morski Instytut Rybacki, Gdynia, Poland. Kubota, M. and Sakai, K. (1978). Autolysis of Antarctic krill protein and its inactivation by combined effects of temperature and pH. Transactions of the Tokyo University of Fisheries, No. 2, 53-63. Kunachowicz, H., Czarnowska-Misztal, E., Klys, W., Wicinska, M. and Jania, M. (1978). Assessment of nutritional value of semiprocessed products of krill. 2. Investigations of the nutritional value of protein. Rocznik Panstwowego Z a k l d u Hygieny, 29, 585-592. Kuznetsova, N. A. (1977). Seasonal distribution of euphausiids in the Kuroshio Current Area. I n “Investigations on Fish Biology and Fishery Oceanography,” Vol. 8, pp. 35-43. TINRO, Vladivostok. Land, M. F. (1978). Animal eyes with mirror optics. Scientific American, 239, 126-134. Land, M. F. and Burton, F. A. (1979). The refractive index gradient in the crystalline cones of the eyes of a euphausiid crustacean. Journal of Experimental Biology, 82, 395-398. Land, M. F., Burton, F. A. and Meyer-Rochow, V. B. (1979). The optical geometry of euphausiid eyes. Journal of Comparative Physiology, 130, 49-62. Lee, 8.4. (1978).Foodand feeding habits of ribboniishes Trichiuruajaponicus and T . lepturua. Bulletin of the Institute of Zoology of the Academia Sinica (Taipei), 17, 117-124. Lestev, A. V. (1978). Krill fishing technology. Rybnogo Khozyaistva, Morskogo, NO. 10, 52-55. Lipskaya, N. J. and Drobysheva, S. S. (1978). Rate of metabolism in some euphausiids of the south Pacific. Transactions of the P. P. Shirshov Institute of Oceanology, 112, 126-132. Lipskaya, N. J. and Drobysheva, S. S. (1979). Rate of oxygen consumption in some euphausiids and hyperiids from the southern Pacific. Abstracts X I V Pacific Science Congress, U S S R , Khabarovsk, August, 1979, p. 63. Lukowski, A. B. (1978). DDT and its metabolites in Antarctic krill (Euphauaia auperba Dana) from South Atlantic. Polskie Archiwum Hydrobiologii, 25, 663-668. McConnaughey, T. and McRoy, C. P. (1979). Food-web structure and the fractionation of carbon isotopes in the Bering Sea. Marine Biology, 53, 257-262. Macpherson, E. (1979). Ecological overlap between macrourids in the western Mediterranean Sea. Marine Biology, 53, 149-159. Makarov, R. R. (1979a). Larval distribution and reproductive ecology of T h y sano&sa mcrura (Crustacea : Euphausiacea) in the Scotia Sea. Marine Biology, 52, 377-386. Makarov, R. R . (1979b). Early larval stages of Antarctic euphausiids (Crustacea: Euphausiacea). Zoologicheskii zhurnal, Moscow, 58, 314-327. Makarov, R . R. (19790). Spawning terms of Antarctic euphausiids. Biologiya Morya, Akademiya Nauk, SSSR, No. 3, 30-38.

620

THE BIOLOGY OF EUPHAUSIIDS

Marshall, N. B. and Merrett, N. R. (1977). The existence of a benthopelagic fauna in the deep sea. I n “A Voyage of Discovery” (Ed. M. Angel), pp. 483497. Pergamon Press, Oxford. Matuda, K., Kanda, K., Koike, A., Takeuchi, S., Inoue, K. and Takasu, K. (1978). A new midwinter trawl for sampling constant depth horizon. Transactione of the Tokyo University of Fisheries, No. 2, 1-25. Matuda, K., Kanda, K., Hamada, E. and Arimoto, T. (1979). On continuous sampling of Antarctic krill. Tranrractions of the Tokyo University of Fisheries, NO. 3, 83-92. Mauchline, J. and Fisher, L. R. (1969). The biology of euphausiids. “Advances in Marine Biology,” Vol. 7, 454 pp. Meyer, T. L., Cooper, R. A. and Langton, R. W. (1979). Relative abundance, behavior, and food habits of the American sand lance, Ammodytes americanua, from the Gulf of Maine. Fishery Bulletin, National Oceanic A t m o q h r i c Adminiatration of the United States, 77, 243-253. Meyer-Rochow, V. B. and Walsh, S. (1978). The eyes of mesopelagic crustaceans. 111. Thysanopoda tricuspidata (Euphausiacea). Cell Tissue Research. 195, 59-79. Mohr, H. (197%). LottechnischeUntersuchungen.Archiv fiir Fischerei~.asenschaft, 30, 85-89. Mohr, H. (197913). Fangtechnische Untersuchungen. Archiv fiir Fiachereihssenschaft, 30, 91-95. Mohr, H. ( 1 9 7 9 ~ ) Fangbeeinflussende . Verhaltensweisen des Krills. Archiv fiir Fischereiwissenschaft, 30, 95-97. Morejohn, G. V., Harvey,’J. T. and Krasnow, L. T. (1979). The importance of Loligo opalescens in the food web of marine vertebrates in Monterey Bay, California. United States Department of Fish and Game, Fish Bulletin, 169, 67-98. Murano, M., Segawa, S. and Kato, M. (1979). Moult and growth of the Antarctic krill in laboratory. Transactions of the Tokyo University of Fisheries, No. 3, 99-106. Musaeva, E. I. (1978). Respiration rates of planktonic animals in the southwestern pert of the Pacific Ocean. Transactions of the P. P. Shirshov Institute of Oceanology, 112, 122-125. Nagayama, F., Yasuike, T., Ikeru,K.and Kawamura, C. (1979). Lipase, mrboxylesterase and catecholoxidase of the Antarctic krill. Transactions of the Tokyo Univeraity of Fisheries, No. 3, 153-159. Nast, F. (1979). Pischereibiologische Untersuchungen am Krill. Archiv f i i r Fkhereiwiesenschaft, 30, 62-67. Naumov, A. G. (1978). The main features of the biology and distribution of Antarctic krill. Rybnogo Khozyaistva, Morslcogo, No. 10, 9-12. Neilson, J. D. and Gillis, D. J. (1979). A note on the stomach contents of adult Atlantic salmon (Salmo salar, Linneaus) from Port Burwell, Northwest Territories. Canadian Journal of Zoology, 57, 1502-1503. Nepgen, C. S. de V. (1979). The food of the snoek, Thyrsites atun. Fisheries Bulletin of South Africa, 11, 39-42. Odate, K. (1978). Distribution of kuril in the northwestern North Pacific. Bulletin of the Tohoku Regional Fisheries Research Laboratory, 39, 21-27. Odate, K. (1979). An euphausiid Crustacea exploited along the Sanriku and Joban coast. Bulletin of the Tohoku Regional Fisheries Research Laboratory, 40, 15-25.

ADDENDUM

621

Okamura, 0. (1970). Studies on the macrouroid fishes of Japan-morphology, ecology and phylogeny. Reportrr of the Usa Marine Biological Station, 17, 1-179. Otero, H. 0. (1977). Length-weight relationship and nutrition o f the “polaca” (Gadidae, Micromeaistius australis Norman, 1937) of the south-west Atlantic. Physis, Seceion A : Los Oceanos y s w O r g a n i m s , 37 (93), 13-23. Pearcy, W. G., Hopkins, C. C. E., Gronvik , S. and Evans, R. A. (1979). Feeding habits of cod, capelin, and herring in Balsfjorden, northern Norway, JulyAugust 1978: the importance of euphausiids. Saraia, 64, 269-277. Pieper, R. E. (1979). Euphausiid distribution and biomass determined acoustically a t 102 kHz. Deep-sea Research, 26, 687-702. Podrazhanskaya, S. G. (1968). Feeding of Macrurus rupestris in the Iceland area. Annalea Biologiques, 1967, 197-198. Pogodin, A. G. (1979). Spawning of euphausiids (Euphausiacea) and duration of development of their younger larvae in waters of northern part of the Tatarsky Strait. Abstracts X I V Pacific Science Congress, USSR, Khabarovsk, August, 1979. 128-129. Ponomareva, L. A. and Drobysheva, S. S. (1978). Euphausiids of the AustralianNew Zealand region and near-by waters of subantarctic. Transactions of the P . P . Shirahov Imtitute of Oceanology, 112, 111-117. Prince, P. A. (1980). The food and feeding ecology of blue petrel (Halobaena caerulea) and dove prion (Pachyptila dmolata).Journal of Zoology, 190, 59-76. Pugh, W. L. (1972). Collections of midwater organisms in the Cariaco Trench, Venezuela. Bulletin of Marine Science, Gu(f of Caribbean, 22,592-600. Rakusa-Suszczewski, S. (1978). Environmental conditions within krill swarms. Polakie Archiwurn Hydrobiologii, 25, 585-587. Rakusa-Suszczewski, S. and Opalinski, K. W. (1978). Oxygen consumption in Euphausia superba. Polakie Archiwum Hydrobiologii, 25, 633-641. Rembiszewski, J. M., Krezeptowski, M. and Linkowski, T. B. (1978). Fishes (Pisces) as by-catch in fisheries o f krill Ewphausia superba Dana (Euphausiacae Crustacea). Polakie A r c h i w r n Hydrobiologii, 25, 677-695. Roschke, N., Flechtenmacher, W. and Schreiber, W. (1978). Extraktionsversuche mit Krillfarcen und Krillkoagulat. Fette Seijen Anstrichmittel Emaehrungsindustrie, 80, 413-416. Rowedder, U. (1979). Feeding ecology of the myctophid Electrona a n t a r c t k (Giinther, 1878) (Teleostei). Meereaforschung, Reports on Marine Research. 27, 252-263.

Saprykina, M. I. (1979). Reproductive cycle of euphausiids in the northern Sea of Japan. Abstracts X I V Pact@ Science Congress, USSR, Khabarovsk, August. 1979, 132-133.

Schreiber, W., Allin, K., Christians, O., Flechtenmacher, W., Oehlenschliiger, J., Papajewski, H., Piske, J., Rehbein, H., Reinacher, E., Wanke, W. and Wilckens, R. (1979). Verarbeitung und Produktentwicklung. Archiv f u r Fkxhereiwiseenschaft, 30, 98-1 12. Secomska, B. and Jelinska, M. (1979). Assessment of the nutritional value of semiprocessed food product obtained from krill. Part 3. Studies in the content of selected vitamins. Rocznik Panatwowego Zakladu Hygieny, 30, 13-18. Segawa, S., Kato, M. and Murano, M. (1979). Oxygen consumption of the Antarctic Krill. Transactiom of the Tokyo University of Fisheries, No. 3, 113- 119.

Shushkina, E. A. and Pavlova, Ye. P. (1973). Metabolism rate and production of zooplankton in the equatorial Pacific. Oceanology, 13, 278-284.

622

THE BIOLOGY OF EUPHAUSIIDS

Shushkina, E. A. and Sokolova, I. A. (1972). Caloric equivalents of the body maw of tropical organisms from the pelagic region of the ocean. Oceanology, 12, 718-724.

Soevik, T. and Braekkan, 0. R. (1979). Fluoride in Antarctic krill (Eupha& superba) and Atlantic krill (Meganyctiphanea norvegica). Journal of the Fisheries Research Board of Canada, 36, 1414-1416. Suzuki, M., Kobayashi, T. and Yanagimoto, M. (1979). Thermal characteristics of Antarctic krill, Euphawia mperba. Bulletin of the Japanese Society of Scientific Fisheries, 45,745-751. Takahashi, T. (1979). Distributions of underwater irradiance, turbidity and suspended matter in the Antarctic Ocean and their relations to Antarctic krill distribution. Transaetions of the Tokyo University of Fisheries, No. 3, 61-70. Tamari, M. (1979). Isolation and identification of ciliatine (2-aminoethylphos. phonic acid) from lipids of edible Antarctic krill, Euphazcsia superba. Agricultural and Biological Chemistry 43, 651-652. Tanaka, K., Matsuda, Y. and Saito, T. (1978). Freezing and defrosting of Antarctic krill Euphauaia superba. Tranaactions of the Tokyo University of Fisheries, No. 2, 65-79. Tanaka, M., Malimoen, A. and Suzuki, K. (19794. Production of canned Antarctic krill. Transactions of the Tokyo University of Fisheries, No. 3, 137-144. Tanaka, K., Matsuda, Y. and Saito, T. (197913). Freezing and defrosting of Antarctic krill. Transactions of the Tokyo University of Fisherim, No, 3, 121-126. Taw, N. and Ritz, D. A. (1979). Influence of subantarctic and subtropical oceanic water on the zooplankton and hydrology of waters adjacent to the Dement River Estuary, south-eastern Tasmania. Azcstralian Journal of Marine and Freshwater Research, 30, 179-202. Tomo, A. P. and Marschoff, E. (1978). Calculo de algunos parametros que hacen a1 mancjo de Euphausia superba Dana, como recurso renovable. Contribucion del Instituto Antarctic0 Argentino, No. 200, 1-19. Toyama, K. and Yano, W. (1979). Application of the juice extractor to the Antarctic krill for the elimination of the shell components. Bulletin of the Japanese Society of Scientific Fisheries, 45,407. Toyama, K., Saito, T. Iwata, Y., Chen, T.-S. and Yano, W. (1979). Preparation and concentration of proteinaceous juice from Antarctic krill. Tramaction8 of the Tokyo University of Fisheries, No. 3, 127-135. Tranter, D. J. and Smith, J. E. (1979). The Antarctic “Biomass” program. Search, 10, 304-307. Valentine, J. W. and Ayala, F. J. (1978). Adaptive strategies in the sea. I n “Marine Organisms, Genetics, Ecology and Evolution” (Eds, B. Battaglia and J. A. Beardmore), pp. 323-345. Plenum Press, New York and London. Worner, F. G. (1979). Zooplankton und Mikroenktonfiinge. Archiv fiir Fischereiwksenschaft, 30, 40-61. Katanabe, E., Suzuki, K., Yagi, T. and Hibikiya, T. (1979). Recovery of the insoluble proteins on the production of soluble concentrates of Antarctic krill. Transactions ofthe Tokyo University of Fisheries, No. 3, 145-151. Williamson, M. H. (1978). The ordinationof incidence data. Journal of Ecology, 66, 911-920. Yanagimoto, M., Kato, N., Yokohama, Y., Kobayashi, T. and Kimura, S. ( 19794. Chemical compositions of Antarctic krill (Euphauaia superba) for the evaluation of processing. Bulletin of the Japanese Society of Scientific Fisheries, 45, 369-374.

ADDENDUM

623

Yanagimoto, M., Yokohama, Y. and Kobayashi, T. (1979b). An application of water jet to doe-viscera methods of Antarctic krill (Euphaueia superba). Bulletin of the Japanese Society of Scientific Fisheries, 45, 375-378. Zelicman, E. A., Lukashevich, I. P. and Drobysheva, S. S. (1978). Aggregative distribution of Thysanoika inemnis (Kroyer) and T . raschii (M. Sam) (Euphausiacea)in the Barents Sea. Okeanolgija, Moscow, 18, 1077-1084. Zelicman, E. A., Lukashevich, I. P. and Drobysheva, S. S. (1979). Round-theyear diurnal vertical migrations of euphausiids Thysanoiksa inerrnis and T . rashii in the Barents Sea. Okeanologija, Moscow, 19, 132-136.

ADDENDUM TO BIOLOGY OF MYSIDS Additional papers have been published on a wide range of topics since the manuscript was submitted for publication. The information in these papers is briefly assessed here under the original chapter numbers and headings so that cross-reference should be facilitated. Gordon (1980) has reviewed the 70-year span of the joint contributions of Walter M. Tattersall and Olive S. Tattersall to research on peracaridan Crustacea and appends a list of their publications.

2. The species of mysids Variation in morphological characters within species is discussed on p. 6. A biometrical study of Spelaeomysis bottazzii by Pesce and Cicolani (1979) demonstrates considerable variation in some of the characters that are used to separate species within t'he genus. Wooldridge (1978) has described two new littoral species from beaches in South Africa: Castrosaccus bispinosa and C. longifissura. A new genus Holmesimysis has been created by Holmquist (1979) to accommodate Acanthomysis costata (Holmes, 1900), Acanthomysis sculpta (W. Tattersall, 1933) and Acanthomysis nuda (Banner, 1948) referred to on p. 11 as Acanthomysis sculpta nuda (Banner, 1948). Two new species, w m e s i m y s i s sculptoides and H . nudensis, are described by Holmquist. All these species occur on the Pacific Coast of North America and confusion has existed between these species and also Acanthomysis davisi Banner, 1948. Consequently, references to the biology of A. davisi, A . costata and A. sculpta prior to the publication of Holmquist's paper must be treated with reserve. A new heteromysinid genus Platymysis is established by Brattegard ( 1980) to accommodate a peculiar dorso-ventrally compressed species that he names P. facilis. This species was collected on the Saba Bank, Caribbean Sea, approximate position 1 7"20'N, 63'33'W, at depths ranging from 16 m to 44 m. Males measured up to 4.1 mm and females up to 3.9 mm in total body length. One female of 3.9 mm had two advanced larvae within the marsupium. Most individuals were collected in the washings from sponges and Brattegard thinks they may spend appreciable periods crawling over the surfaces of sponges in areas in which the sea is in constant motion. 624

ADDENDUM

625

Key to Genera The following should be inserted in the “Key to Genera” on p. 35 in place of the last paragraph of Item 86: Exopod of pleopod IV of two segments, terminating distally in two short, spiny, peg-like structures .. .. .. Holme&mysis Exopod of pleopod IV of two segments, terminating distally in two setae (Fig. 9.16) .. .. .. .. .. .. . . 87 The following should be inserted in the Key to Genera on p. 36, between sections 89 and 90: Body dorso-ventrally compressed; abdominal somites with welldeveloped pleural plates ., .. .. .. . . Platymysis Body not dorso-ventrally compressed; abdominal somites without distinct pleural plates . . .. .. .. .. .. . . 90

3. The larvae in the marsupium w e a s e of young from the marsupium is discussed on p. 65. Amaratunga and Corey (1979) found that young of Mysis stenolepis are released singly at intervals of 3-10 min, the entire brood being released over a period of 36 hours. Actions of the female, such as rhythmic expansion of the marsupium and flexing of the abdomen, act in conjunction with movements of the young to effect release to the environment. Modlin (1979) defines four recognizable embryonic stages in this species similar to those described on p. 59. Ladurantaye and Lacroix (1980) found that the brood size of Mysis litoralis was 33-113 and seemed to be related to the body length of the female in Saguenay Fjord at 70”N off the Gulf of St Lawrence. The body length of ovigerous females ranged from 15-5 mm to 25 mm. The duration of development of the larvae within the marsupium is about 6 months, the larvae having a total length of 3.8 mm on emergence. The numbers of young carried by ~ a p h r o m ~ bs ~ om s a n i in the marsupium range from 4 to 9 in individuals of 6.7-10-0 mm body length from Texas (27”36’N, 97’18‘W) (Compton and Price, 1979).

4. Vertical distribution and migration Equipment and nets for sampling mysids are discussed on p. 66. Brunel et al. (1978) have published a description of enlarged versions

626

THE BIOLOGY OF MYSIDS

of the Macer sledge-mounted suprabenthic sampler which are suitable for examining micrix3tratification of species above the sediment. Many species of rqyids live in close association with the sediment, some even within it during the day, but move upwards into the water column at night. Hobson and Chess (1979) designed a meroplankton trap in which they found unidentified mysids that had emerged from the lagoon floors of Kure and Midway Atolls, Hawaii at night. The possibility of detecting mysid populations through use of sonar equipment is briefly discussed on p. 69. Jukic (1978) suggests that Lophogaster typicus in conjunction with two species of euphausiids was responsible for a sonic scattering layer detected with a 30 kHz sounder in the Adriatic Sea. Brunel (1979) has examined seasonal changes in the diel vertical migrations of Erythrops erythrophthalma, Meterythrops robusta, Mysis mixta and Pseudomma afine in the Gulf of St Lawrence. Irregularities were present in the patterns of the migrations seasonally and between different years. Seasonal changes in the vertical distribution of My& Zitoralis in Saguenay Fjord are described by Ladurantaye and Lacroix ( 1 980).

5 . Food and feeding Fluorescence of the gut contents of the freshwater Mysis relicta was used to determine the relative quantities of phaeopigments present by Grossnickle (1979). The results indicate that this species filters phytoplankton throughout the summer diel vertical migrations in Lake Michigan at night. A method using the fluorescent dye, Rhodamine b, to determine predation rates of M . relicta on copepods has been developed by Lasenby (1979) for use in laboratory and field situations. An unstained population of a known number of copepods is innoculated with a known number of stained individuals. Mysids then graze the copepods. The number of stained survivors at the end of the experiment can be quickly counted and converted to the total number of copepods consumed. A single M . relicta was found to consume 0.15-1.35 Diaptomus ashlandi per hour in Kootenay Lake, British Columbia and 3.2 Limnocalanus macrurus per hour on Stony Lake, southern Ontario. Odum and Heald (1972) examined the gut contents of 120 Taphromysis bowmani and found the following by volume: 9% diatoms, 7% copepod fragments, 26% fine vascular plant detritus, 39% inorganic material and 19% unidentified material and particles. Their comparable analyses of the food of 153 Mysidopsis almyra is: 6% diatoms, 5% copepod fragments, 31% fine vascular plant detritus,

ADDENDUM

627

32 yo inorganic material and 26% unidentified material and particles. The popufations studied were from a mangrove community in Florida. Smaller Mysis relicta preyed on smaller organisms than larger M . relicta and showed preferences for certain prey species in experiments made by Cooper and Goldman (1980). They attributed patterns of prey selection to the mechanical efficiency with which the M . relicta captured and handled the different sized particles, to the ability of the prey organism to escape and to predator-prey encounter frequencies.

6. Chemical composition Dry body weight is 0.170 wet weight in Mysis caspia; that is, the water content is 83% (Bondarenko, 1978b). The carbon : nitrogen ratio in Neomysis rayii is 4.1 according to McConnaughey and McRoy (1979); this is within the wide range shown in Table XI1 (p. 101). The relationship between the calorific equivalent of body weight, W e in cal/individual, and body length in mm in tropical mysids, the species of which were not named, was determined by Shushkina and Sokolova (1972)as: W e = 0.024 La*''

7. Internal anatomy Osmotic regulation of the blood concentrations of mysids is described on p. 121. McLusky (1979), examining Praunus flexuosus from the Isefjord in Denmark, found the isosmotic point to be 21-22%, at 5°C and 15°C. He determined the ionic composition of the blood in terms of sodium, potassium, magnesium, calcium and chloride and the changes ~ . relative ionic taking place in it at salinities of 10, 18 and 2 7 ~ 5 %The composition of the blood of P . jIexu0su-s is compared with that of Leptomysis mditerranea and other crustaceans. The blood concentrations of the ions, when totalled, accounted for the osmotic concentrations. The blood concentrations of Praunus flexwsus from the Isef jord, however, were significantly lower than those found in the same species from western Scotland, a phenomenon considered to indicate a long-term acclimation.

8. Physiology and responses to physical-chemical parameters of the environment

A study of the effects on the rate of respiration of Neomysis mirabilis

628

THE BIOLOGY OF MYSIDS

in different sized respirometers containing different densities of mysids was made by Kuz’micheva &nd Kukina (1974) with negative results. Shushkina and Pavlova (1973) related the metabolic rate T,expressed as m cal/day/individual, to the body weight M , expressed as m call individual in tropical mysids that they did not identify. They found the following relationship :

T = 13.3 MO.498 Bondarenko (1978b) examined the respiration rates of Mysis caspia and other Caspian Sea species. Regression equations are given along with a review of comparable equations from the literature. The littoral Gastrosaccus psammodytes exhibited a rhythm of oxygen consumption, high levels coinciding with high tide, low levels with low tide. The rhythm was independent of light intensity (Dye, 1980). The rate of oxygen consumption was more sensitive to temperature in winter than summer but showed a greater dependence on body size in summer than in winter. Respiration, oxygen consumption and carbon dioxide production were studied in Mysis relicta by Ranta and Hakala (1978) whose results conform with those previously published on this species. The possibility that the mitochondria1 and microsomal electron transport system (ETS) may provide an indirect method of measuring respiration rate is discussed briefly on p. 150. Bamstedt (1979), however, in studying a range of planktonic organisms including Boreomysis arctica and Hernimysis abyssicola found that ETS and respiratory rate varied seasonally but that the variations were poorly correlated. He concludes, therefore, that ETS may not be a practical method of predicting population metabolism. Behavioural reactions of mysids to environmental illumination are discussed on p. 152. Macquart-Moulin (1979) has compared the behaviour of a hypogean, that is cave-dwelling, species Hemimysis speluncola with that of the littoral Leptomysis lingvura. The former species exhibits strong negative phototaxis above l o - * lux and a clear positive reaction at 10-41ux while L. lingvura shows only slight orientation. The reactions of Hemimysis spelunwla are similar t o those of species burrowing deeply into sediment or residing under rocks during the day. The salinity tolerance of Praunus jlexuosus in western Scotland is defined on p. 154. McLusky (1979) examined this aspect of P . jlexuosw in the Isefjord, Denmark. Winter animals maintained at low temperature showed enhanced survival at low salinities over summer animals kept in the same conditions.

ADDENDUM

629

9. Behaviour The burrowing behaviour of species in the genus Gaetrosaccus is discussed on p. 185 where it is stated that there is little conclusive evidence of burrowing of species in other genera. Kamihira (1979), however, has examined the distribution of Archaeomysis kokuboi on a sandy beach relative to its ability to burrow and withstand desiccation.

10. Population dynamics

A biometrical study of Spelaeomysis bottazzii is described by Peace and Cicolani (1979) who examine the relationships between body length on the one hand and the numbers of spines on the exopod of the uropod, the proximal width of the telson, and the antennal scale length on the other. Regression equations are given for these relationships and for antennal scale width on length. An account of the life cyde of Mysis stenozepis is given by Modling (1979) while Ladurantaye and Lacroix (1980) describe the annual cycle of M . litoralis in Saguenay Fjord. This latter species matures to breed at one year old but a few individuals may survive to the following year. Populations of Taphromysis bowmani probably breed throughout the year at Four Bluff, Texas (27’36’N, 97’18’W) (Compton and Price, 1979).

11. Geographical distribution Hypogean species are listed in Table XXXVIII (p. 224). Peace (1976) briefly reviews their number and geographical distribution. The Caspian Sea. Bondarenko (1978a) has examined the distribution of the majority of species listed on p. 232. He does not mention Amnthomysis strauchi, Paramysis bakuensis or Schistomysis elegans. The following four species dominated the mysid fauna of the northern region of the Sea: Paramysis baeri, P . intermedia, P . Eacustris and P . ullskyi. Quantitative distribution maps of their occurrence show that their centres of distribution tend to be segregated and related to depth, substratum and salinity. There has been a significant decrease in the biomass of mysids present in this region which Bondarenko ascribes to the reduction in river discharges that has taken place and which has altered the quantity of suspended material entering the region.

12. Predators and parasites The available information on predation of fish on offshore species of

630

THE BIOLOGY OF MYSIDS

mysids is discussed on p. 238. Some lantern fishes of the family Myctophidae are reported by Gorelova '(1974) to feed on mysids to a small extent. Notothenia microlepidota Hutton eats some mysids off Chile (Pequeno, 1976) as does the southern blue whiting Micromesistius australis Norman in the southern Patagonian region (Otero, 1977). The macrourid Macrourus berglax Lacepede includes mysids as a minor component of its diet in the North Atlantic (Geistdoerfer, 1979). Boreomysis megalops and other unidentified mysids were found by Macpherson (1979) in the food of Hymenocephalus italicus Giglioli, Nezumia aeqwzlis (Gunther), Coelorhynchus coelorhynchus (Risso) and Trachyrhynchus trachyrhynchus (Risso). Haedrich and Polloni (1976) recorded mysids in the stomachs of Coryphuenoides carapinus Goode and Bean as a sub-dominant item. CTnathaphausia zoea occurred in the stomachs of Alepocephalus bairdii (Goode and Bean) while unidentified species also occurred in the food of Hoplostethus mediterraneus (Cuvier) and Glyptocephalus cynoglossus (L.) according to Buit (1978). Gnathophausia zoea is also recorded by Marshall and Merrett (1977) in the stomachs of Alepocephalus rostratus Risso and Bathygadus melanobranchus Vaillant off north-west Africa. The following additional information on predation of mysids by inshore fish in various geographical regions is available. North Atlantic, between about 60"N and 40"N. The megrim, Lepidorhombus whifiagunis (Walbaum), includes Schistomysis ornata as a minor component of its diet in Icelandic waters (Steinarsson, 1979). The American sand lance, Ammodytes americanus De Kay, eats mysids that Meyer et al. (1979) did not identify, occasionally in the Gulf of Maine. Atlantic Ocean, between 40"N and 40"s. Odum and Heald (1972) studied the food of fish within a mangrove community in Florida, USA. The following is a list of species predatory on Taphromysis bowmani, the commoner species, and on Mysidopsis almyra. The percentage of the diet, by volume, represented by mysids is given where available: Scaled sardines, Harengula pensacolae Goode and Bean, 16 % Bay anchovy, Anchoa mitchilli (Valenciennes),19% Sea catfish, Arius felis (L.),7% Rainwater killifish, Lucania parva (Baird), 19% Crevalle jack, Caranx hippos (L.) Silver jenny, Eucinostomus gula (Quoy and Gaimard), &6y0 Spotfin mojarra, Eucinostomus argenteus Baird and Girard, 7-974 Striped mojarra, Diapterus plumieri (Cuvier), 44% Red drum, Sciaenops ocellata (L.) Spotted seatrout, Cynoscion nebulosus (Cuvier)

ADDENDUM

631

Silver perch, Bairdiella chrysura (Lacepixie) Pinfish, Lagodon rhomboides (L.) Crested goby, Lophogobius cyprinoides (Pallas) Code goby, Gobiosoma robustum Ginsburg Clown goby, Microgobius gulosus Tidewater silverside, Menidia beryllina (Cope): mysids are not important in its diet during the day but represent over 60% of its food at night Lined sole, Achirus lineatus (L.) Gulf toadfish, Opsanus beta (Goode and Bean) Off South Africa, Nepgen (1979) notes that the snoek, Thyrsites atun (Euphrasen), occasionally eats Mysidopsis major. Mediterranean. The hake, Merluccius merluccius L., and the horse mackerel, Trachurus trachurus L., include Lophogaster typicus in their diets which consisted primarily of euphausiids, at least during the day (Jukic, 1978). Aral Sea. Predation of mysids by fish in the Aral Sea is discussed by Proskurina (1978). North Paci$c, north of about 40"N. The walleye pollack, Theragra chulcogramma (Pallas), includes unidentified mysids as a very minor component of its diet in the eastern Bering Sea (Bailey and Dunn, 1979).

I n Chilka Lake, India, fingerling Lutes culcarifer (Bloch) feed predominantly on mysids (Patnaik and Jera, 1978). Birds predate on mysids, as discussed on p. 249. The Antarctic mysid, Antarctomysis maxima, is a component of the diets of the burrowing petrels, Halobaena caerulea Gmelin and Pachyptilu cksolata (Gmelin),in South Georgia (Prince, 1980). Several invertebrates prey on mysids (p. 250). In the laboratory, Nelson et al. (1979) used Neomysis mercedis as a food for the shrimp Crangon franciscorum Stimpson. This shrimp and Palaemon macrodactylus Rathbun eat large quantities of Neomysis mercedis in the Sacramento-San Joaquin estuary in central California. Sitts and Knight (1979) estimate predation rates by the populations of these shrimps on the standing crop of N . mercedis. Mysids are parasitized by a wide range of organisms (p. 251). The occurrence of the leech, Mysidobdella borealis (ex. Ichthyobdella borealis Johansson, 1899), on N e m y s i s americana is reported by Burreson and Allen (1978). It occurs most commonly on N . americana along the eastern coast of the United States in the winter and early spring. They also found it attached to Mysis stenolepis in Passamaquoddy Bay but in this species it was most common in the spring and autumn. This

632

THE BIOLOGY OF MYSIDS

leech does not appear to deposit its cocoons in the mysids but may feed on the mysids.

13, Mysids in the marine economy The effects of introducing mysids to various lakes to supplement the food of resident or introduced fish is discussed on p. 261. Morgan et al. (1978) and Goldman et al. (1979) discuss the impact of introduced Mysis re2icta in Lake Tahoe, California, on populations of the three resident cladoceran species that declined as mysid abundance increased.

APPENDIXI. TAXONOMIC LIST OF SPECIES The following new species should be added t o the list on p. 325. Sub-family GASTROSACCINAE Genus Gastrosaccw Norman, 1868 C. bispinosa Wooldridge, 1978 G. longi$ssura Wooldridge, 1978 The following species should be deleted from the genus Acanthomysis Czerniavsky, 1882 on p. 335: A. costata (Holmes, 1900), A. sculpta (W. Tattersall, 1933). The following new genus and list of species should be inserted on p. 337 between the genera Hemimysis and Idiomysis: Genus Holmesimysis Holmquist, 1979 H . costata (Holmes, 1900) H . nuda (Banner, 1948) H . nudensis Holmquist, 1979 H . sculpta (Banner, 1948) H . sculptoides Holmquist, 1979 The following new genus and species should be inserted in the Tribe Heteromysini on pp. 340-341 : Genus Platymysis Brattegard, 1980 P.facilis Brattegard, 1980

APPENDIX11. CLASSIFIEDLIST OF LITERATURE FOR EACHGENUS The following are additional references for inclusion in this Appendix. Acanthontysis: Bondarenko, 1978a; Holmquist, 1979 Anttirctomysis: Prince, 1980

ADDENDUM

633

Antromysis: Pesce, 1976 Archaeomysis: Kamihira,' 1979 Boreomysis: Bamstedt, 1979 Bowmaniella: Cooley, 1978 Caspiomysis: Bondarenko, 1978a Diamysis: Bondarenko, 1978a Doxomysis: Young and Wadley, 1979 Erythrops: Brunel, 1979 Gastrosaccw: Odum and Heald, 1972; Rasmussen, 1973; Wooldridge, 1978; Dexter, 1979; Young and Wadley, 1979 Hemimysis: Bondarenko, 1978a; Bamstedt, 1979 Hetermysis: Brattegard, 1980 Heteromysoides: Pesce, 1976 Holmesimysis: Holmquist, 1979 Katamysis: Bondarenko, 1978a Lirnnomysis: Bondarenko, 1978a Lophgaster: Jukic, 1978 Mesopodopsis: Rasmussen, 1973 Metamysidopsis: Dexter, 1979 Meterythrops: Brunel, 1979 Mysidopsis: Odum and Heald, 1972; Cooley, 1978; Compton and Price, 1979; Nepgen, 1979 Mysis: Bondarenko, 1978a, b; Morgan et al., 1978; Ranta and Hakala, 1978; Amaratunga and Corey, 1979; Brunel, 1979; Grossnickle, 1979; Lasenby, 1979; Cooper and Goldman, 1980; Ladurantaye and Lacroix, 1980 Neomysis: Rasmussen, 1973; Kuz'micheva and Kukina, 1974; Holmquist, 1979; McConnaughey and McRoy, 1979; Nelson et al., 1979; Orsi et al., 1979; Sitts and Knight, 1979 Paramysis: Bondarenko, 1978a Platymysis: Brattegard, 1980 Praunw: Rasmussen, 1973; McLusky, 1979 Pseudomma: Brunel, 1979 Rhopalophthalrnus: Young and Wadley, 1979 Schistomysis: Rasmussen, 1973; Steinarsson, 1979 Siriella: Young and Wadley, 1979 Spelaeomysis: Pesce, 1976; Sbordoni et al., 1978; Pesce and Cicolani, 1979 Stygiomysis: Pesce, 1976 Taphromysis: Odum and Heald, 1972; Cooley, 1978; Compton and Price, 1979 Troglomysis: Pesce, 1976

634

THE BIOLOGY OF MYSIDS

APPENDIX111. CLASSIFIEDLIST OF LITERATURE FOR GEOGRAP~~ICAL ' REGIONS The following additional references should be included in this Appendix. Freshwater environments: Pesce, 1976; Morgan et al., 1978; Grossnickle, 1979; Lasenby, 1979 North AtZantic, north of 40"N, excluding the Baltic Sea: Bamstedt, 1979; Brunel, 1979; Steinarsson, 1979; Ladurantaye and Lacroix, 1980 The BalticSea: Rasmussen, 1973; McLusky, 1979 The Western Atlantic, between 40"N and 40"s: Odum and Heald, 1972; Cooley, 1978; Compton and Price, 1979; Dexter, 1979; Brattegard, 1980

The Eastern Atlantic, between 40"N and 40°S,but excluding the Mediterranean Sea, and including South Africa: Wooldridge, 1978; Nepgen, 1979

The MediterraneanSea: Jukic, 1978; Pesce and Cicolani, 1979 The Caspian Sea: Bondarenko, 1978a The Aral Sea: Proskurina, 1978 North Pacijic: Holmquist, 1979 The Eastern PaciJic between 40"N and 40"s:Dexter, 1979; Holmquist, 1979; Orsi et al., 1979; Sitts and Knight, 1979 Japanese Coastal Waters, Sea of Japan, China Seas and the Amtralasian Region: Kamihira, 1979 REFERENCES Amaratunga, T. and Corey, S. (1979). Marsupium and release of young in My& stenolepis Smith (Mysidacea).Cruataceanu,37,8684. Bailey, K. and Dunn, J. (1979). Spring and summer food of walleye pollock, Theragra chulcogramma, in the eastern Bering Sea. Fishery Bulletin, N a t i o d Oceanic Atmospheric Administration of the United States, 77,304-308. Bamstedt, U. (1979). Seasonal variation in the respiratory rate and ETS activity of deep-water zooplankton from the Swedish west coast. I n "Cyclic Phenomena in Marine Plants and Animals" (Eds E. Naylor and R. G. Hartnoll), pp. 267-274. Pergamon Press, Oxford. Bondarenko, M. V. (1978a). The composition and distribution of mysids from the north Caspian Sea. Proceedings All- Union Research InatitUte of Marine Fisheriesand Oceanography ( V N I R O ) ,132 (3), 13-25. Bondarenko, M. V. (1978b). The metabolic intensity in some mysids from the Caspian Sea. Proceedings All- Union Research Institute of Marine P+heries and Oceanography (VNIRO),132 (3), 32-39. Brattegard, T. (1980). Platynaysis facilis Gen. et Sp. nov. (Crustacea : Mysidacea : Hetermomysini) from the Saba Bank, Caribbean Sea. Sarah, 65,49-52.

ADDENDUM

635

Brunel, P. (1979).Seasonal changes of daily vertical migrations in a suprabenthic cold-layer shelf commufiity over mud in the Gulf of St. Lawrence. I n “Cyclic Phenomena in Marine Plants and Animals” (Eds E. Naylor and R. G. Hartnoll),pp. 383-390. Pergamon Press, Oxford. Brunel, P., Besner, M., Messier, D., Poirier, L., Granger, D. and Weinstein, M. ( 1978). Le traineau suprabenthique Macer-GIRO&: appareil am6liorB pour l’echantillonnage quantitatif 6tag6 de la petite f a w e nageuse au voisinage du fond. Internationale Revue der gesamten Hydrobiologie u. Hydrographie, 63, 815-829. Buit, M.-H. du (1978).Alimentation de quelques poissons teleosteens de profondeur dans la zone seuil de Wyville Thomson. Oceanologia Actu, 1,129-134. Burreson, E.M. and Allen, D. M. (1978).Morphology and biology of Mysidobdella borealis (Johansson) comb. n. (Hirudinea :Piscicolidae), from mysids in the western North Atlantic. Journal of Parmitology, 64,1082-1091. Compton, C. E. and Price, W. W.(1979).Range extension to Texas for Taphromysis bowmani Bacescu (Crustacea : Mysidacea) with notes on its ecology and generic distribution. Contributions to Marine Science, 22,121-125. Cooley, N. R. (1978).An inventory of the estuarine fauna in the vicinity of Pensacola, Florida. Florida Marine Research Publications, 31,l-119. Cooper, S . D. and Goldman, C. R. (1980).Opossum shrimp (Mysis relicta) predation on zooplankton. Canadian Journal of Fisheries and Aquatic Sciences, 37,909-919. Dexter, D. M. (1979).Community structure and seasonal variation in intertidal Panamanian sandy beaches. Estuarine and Cowtul Marine Science, 9,543-558. Dye, A. H. (1980).Aspects of the respiratory physiology of Gastrosaccus psammodytes Tattersall (Crustacea : Mysidacea). Comparative Biochemistry and Physiology, 65A,187-191. Geistdoerfer, P. (1979). RBcherches sur l’alimentation de Macroums berglax LacepMe 1801 (Macrouridae, Gadifonnes). A n d e s Z’lnstitut ocianographique, 55,135-144. Goldman, C. R., Morgan, M. D., Threlkeld, T. and Angeli, N. (1979).A population dynamics analysis of the cladoceran disappearance from Lake Tahoe, California-Nevada. Limnology and Oceanography, 24,289-297. Gordon, I. (1980).Walter M. Tattersall and Olive S. Tattersall: 7 decades of peracaridan research. Crustaceans, 38,311-320. Gorelova, T. A. (1974).Zooplankton from the stomachs of juvenile lantern fish of the family Myctophidae. Oceanology, 14,575-580. Grossnickle, N. E. (1979).Nocturnal feeding patterns of Mysia relictu in Lake Michigan, based on gut content fluorescence. Ikmnology and Oceanography, 24,777-780. Haedrich, R. L. and Polloni, P. T. (1976).A contribution to the life history of a small rattail fish, Coryphaenoides uzrapinus. Bulletin of the Southern Cal.liforn& Academy ofSciences, 75,203-211. Hobson, E. S. and Chess, J. R. (1979).Zooplankters that emerge from the lagoon floor at night a t Jure and Midway Atolls, Hawaii. Fishery Bulletin, National Oceanic Atmospheric-Administration of the United States, 77,275-280. Holmquist, C. (1979).Mysis costata Holmes, 1900 and its relations (Crustacea, Mysidacea). Zoologische Jahrbucher, Systematik, Okoligie und Cfeographie der Tiere, 106,471-499. Jukic, S. (1978). Contribution to the knowledge of the relationship between

636

THE BIOLOGY OF MYSIDS

formation of deep scattering layer (DSL) and biological components in the central Adriatic. Acta Adriatica, 19,3-15. Kamihira, Y. (1979). Ecological studies of macrofauna on a sandy beach of Hakodate, Japan. 11. On the distribution of peracarids and the factors influencing their distribution. Bulletin of the Faculty of Fisheries, Hokkaido Uniuersity, 30, 133-143. Kuz’micheva, V. I. and Kukina, I. V. (1974). Intensity of respiration of mysids when packed in respirometers a t different intensities. Oceanology, 14, 732-736. Ladurantaye, R. de and Lacroix, G. (1980). RBpartition spatiale, cycle saisonnier et croissance de Mysis litoralis (Banner, 1948) (Mysidacea) dans un fjord subarctique. Canadian Journd of Zoology, 58,693-700. Lasenby, D. C. (1979). A method for determining predation rates of macrozooplankton. C a d i a n Journal of Zoology, 57,1504-1506. McConnaughey, T. and McRoy, C. P. (1979). Food-web structure and the fractionation of carbon isotopes in the Bering Sea. Marine Biology, 53,257-262. McLusky, D. S. (1979). Some effects of salinity and temperature on the osmotic and ionic regulation of Praunw jlexuosus (Crustacea, Mysidacea) from Isefjord. Ophelia, 18, 191-203. Macpherson, E. (1979). Ecological overlap between macrourids in the western Mediterranean Sea. Marine Biology, 58,149-159. Macquart-Moulin, C. (1979). Ecophysiologie comparee des mysidacds Hemimysis speluncola Ledoyer (cavernicole) et Leptomysis lingvura G. 0 . Sars (noncavernicole). L’orientation B la lumikre: tests ponctuels. Journd of Experimental Marine Biology and Ecology, 38,287-299. Marshall, N. B. and Merrett, N. R. (1977). The existence of a benthopelagic fauna in the deep sea. I n “A Voyage of Discovery” (Ed. M. Angel), pp. 483-497. Pergamon Press, Oxford. Meyer, T. L., Cooper, R. A. and Langton, R. W. (1979). Relative abundance, behaviour and food habits of the American sand lance, Ammodytes americanus, from the Gulf of Maine. Fishery Bulletin, National Oceanic Atmospheric Administration of the United States, 77, 243-253. Modlin, R. F. (1979). Development of Mysis stenolepis (Crustacea : Mysidacea). American Midland Naturalist, 101,250-254. Morgan, M. D., Threlkeld, S. T. and Goldman, C. R. (1978). Impact of the introduction of kokanee (Oncorhynchus nerka) and opossum shrimp ( M y s k relicta) on a subalpine lake. Journal of the Fwheries Research Board of Canada, 35,1572-1579.

Nelson, S . G., Simmons, M. A. and Knight, A. W. (1979). Ammonia excretion by the benthic estuarine shrimp Crangonfranciscorum (Crustacea : Crangonidae) in relation to diet. Marine Biology, 54, 25-31. Nepgen, C. S. de V. (1979). The food of the snoek, Thyrsites atun. Fisheries Bulletin ofSouth Africa, 11,39-42. Odum, W. E. and Heald, E. J. (1972). Trophic analyses of an estuarine mangrove community. Bulletin of Marine Science, Gulf and Caribbean, 22,671-738. Orsi, J. J., Knutson, A. C. and Fast, A. W. (1979). An extension of the known range of Neomysis mercedis, the opossum shrimp. Cali,fornia F k h and Game, 65,127-130.

Otero, H. 0. (1977). Length-weight relationship and nutrition of the ‘‘polaca” (Gadidae, Micromesistius australis Norman, 1837), of the south-west Atlantic. Physis, Seccion A : Los Oceanos y Sus Organkmos, 37 (93), 13-23.

ADDENDUM

637

Patnaik, S. and Jena, S. (1978). Some aspects of biology of Lutes calcarijer (Bloch) from Chilka Lake. lhdian Journal of Fisheries, 23 (1975), 65-71. Fequeno, G. R. (1976). New antecedents for Notothenia microlepidota Hutton (Teleostomi, Nototheniidae). Noticiario Mensual Museo Nmional de Historia Natural (Santiago), 21 (241), 5-8 Pesce, G. L. (1976). Stato attuale delle conoscenze sui Misidacei cavernicole e freatici (Crustacea). Notiziario del C ~ T C O Speleologica ~O Romano, 1, 47-57. Pesce, G. L. and Cicolani, B. (1979). Variation of some diagnostic characters in Spelaeomysis bottazzii Caroli (Mysidacea).Crustaceana, 36,74-80. Prince, P. A. (1980). The food and feeding ecology of blue petrel (Halobeana caerulea) and dove prion (Pachyptila desolata). Journal of Zoology, 190, 59-76. Proskurina, E. S. (1978). Nutrition of juvenile fish in the Aral Sea. Voprosy Ikhtiologii, 18,46&466. Ranta, E. and Hakala, I. (1978). Respiration of Mysis relicta (Crustacea, Malacostraca). Archiv f u r Hydrobiologie, 83, 515-523. Rasmussen, E. (1973). Systematics and ecology of the Isefjord marine fauna (Denmark) with a survey of the eelgrass (Zostera) vegetation and its communities. Ophelia, 11, 1-507. Sbordini, V., Pesce, L., Colognola, It. and Fusacchia, F. (1978).Differenziamento genetic0 tra popolazioni du Spelaeomysis bottazzii (Crustacea, Mysidacea). I n “Biogeografia delle Caverne Italiane”, pp. 85-86. X X I I Congress0 Nazionale Societa Italiana di Biogeografia, Verona-Costagrande, 15-18 Giugno, 1978. Shushkina, E. A. and Pavlova, Ye. P. (1973). Metabolism rate and production of zooplankton in the equatorial Pacific. Oceanology, 13, 278-284. Shushkina, E. A. and Sokolova, I. A. (1972). Calorie equivalents of the body mass of tropical organisms from the pelagic region of the ocean. Oceanology, 12, 718-724.

Sitts, R. M. and Knight, A. W. (1979). Predation by the estuarine shrimps Crangon franciscorum Stimpson and Palaemon macrodactylus Rathbun. Biological Bulletin, Woods Hole Marine Biological Laboratory, 156, 356-368. Steinarsson, B. (1979). The food of lemon sole (Microstomus kitt Walbaum), megrim (Lepidorhombus whifiagonis Walbaum) and witch (Glyptocephalus cynogloasus L.) in Icelandic waters. Meeresforschung, Reports on Marine Research, 27, 156-171. Wooldridge, T. (1978). Two new species of Gastrosaccus (Crustacea, Mysidacea) from sandy beaches in Transkei. Annals of the South African Museum, 76, 309-327.

Young, P. C. and Wadley, V. A. (1979). Distribution of shallow-water epibenthic macrofauna in Moreton Bay, Queensland, Australia. Marine Biology, 53, 83-97.

This Page Intentionally Left Blank

Taxonomic Index A 178, 181, 184,233,248, 336, 624, Abudefduf 632 leucopomus, 248 sculpta nuda, 11, 624 Acanthephyra sinensis, 336 purpurea, 534 stelleri, 336 Acanthocephrtla, 541 strauchi, 48, 190, 232, 255, 336, 629 Acanthomysis, 35, 84, 134-136, 140, tamurai, 336 179,184,233,335,343,632 trophopristes, 336 alaskensis, 335 Achirus anomala, 335 lineatw, 631 aokii, 335 Acineta aspera, 335 tuberosa, 253 borealis, 335 Acipenser columbiae, 335 medirostris, 247 costata, 233, 335, 624, 632 tranamontanus, 241 davisi, 335, 624 Aethia dimorpha, 335 crisiWella, 249, 535 dybowskii, 335 pusilla, 249, 636 fujinagai, 335 Afromysis, 16, 30, 31, 88, 331, 343 grimmi, 232, 335 bainbridgei, 33 1 hodgarti, 335 dentisinus, 33 1 hwanhaienais, 335 guinensis, 48, 331 indica, 335 hansoni, 332 inJEata, 232, 335 mawopsia, 332 japonica, 335 ornata, 332 koreana, 335 Agonus longkornis, 104, 115, 335 cataphractus, 241 longicornis glabra, 11 Aldrovandia, 239 longirostris, 335 Alepocephalus macrops, 335 bairdii, 630 macropsis, 69, 233, 335 rostratus, 630 mitaukurii, 261, 335 Allothunnus nakazatoi, 336 fallai, 530 nephrophthalma, 336 Alosa okayamaensis, 336 pseudoharengus, 243 orrzata, 336 sapidissimu, 241 p e h g i m , 336 Amathimysia, 25, 84, 172, 326, 343 platycauda, 336 cherados, 48, 172, 326 pseudomacropsis, 221, 233, 254, 336 gibba, 48, 172, 326 pseudomitsukurii, 336 polita, 48, 172, 326 q d r i s p i n o s a , 336 Amblyops, 21, 123, 326, 344 sagamiensis, 336 abbrevkta, 48, 129, 130, 184, 220, schrencki, 336 227, 229, 256, 326 aculpb, 48, 51, 61, 6 4 , 7 3 , 9 5 , 9 6 , 141, aequispina, 69, 326 639

640

TAXONOMIC INDEX

antarctica, 235, 326 durbani, 326 ewingi, 234, 327 kempi, 104, 105, 114, 229, 327 magna, 3, 69, 157, 327 spinifera, 327 tattersalli, 235, 327 tenukxuda, 327 trisetosa, 327 Amblyopsoides, 27, 123, 327, 344 crozetii, 235, 327 obtusa, 327 ohlinii, 227, 327 Ammodytes arnericunus, 243, 609, 630 Amphidinium klebsii, 458 Amphilina foliacea, 255 Amphipoda, 4 Anaplopoma Jimbria, 610 A w h d i n a , 20, 88, 90, 324, 344 agilis, 43, 45, 48, 61, 71-74, 91, 109, 110, 114, 141, 152, 161, 166, 171, 176, 181, 182, 194, 245, 250,254, 324 d a n b n i , 324 dentata, 324 flemingi, 325 grossa, 325 latifrons, 325 madagascariensis, 7, 325 media, 325 obtusifrons, 325 oculata, 48, 231, 325 penicillata, 325 sanzoi, 7, 231, 325 truncata, 174, 325 typica, 48, 73, 74, 172, 226, 325 typica orientalis, 11 zimmeri, 325 Awhoa mitchdli, 630 mitchilli diaphana, 244 Anisakis sp., 329, 540, 541 Anisomysis, 33, 36, 128, 336, 344 aikawai, 336 bqurcuta, 336 bipartoculata, 336 hanseni, 48, 336

hispida, 336 ijirnai, 336 ijimai estafricana, 11 incisa, 336 kunduchiana, 336 (Anisomysis) kunduchiana, 11 lamellicauda, 336 laticauda, 336 lewi, 235, 336 rnaris rubri, 184, 336 megalops, 336 mixta, 336 mixta australis, 11 (Paranisomysis) maris rubri, 11 pelewensia, 336 sirielloides, 184, 336 tattersdlae, 336 thurneysseni, 336 vasseuri, 224, 336 Anophelina, 12 Anotopterus pharao, 532 Antarctomysis, 36, 84, 336, 344, 632 maxima, 48, 62, 83, 84, 86-88, 96, 174, 199, 200, 235, 238, 249, 254, 336,631 ohlinii, 174,235, 238,336 Antonogadus rnegalokynodon, 246, 529 Antromysia, 12, 14, 33, 36, 128, 336, 344, 633 almyra, 223, 336 americana, 337 anophelinae, 48, 186, 233, 337 (Anophelina) anophelinae, 11 baharnenais, 48, 172, 337 cenotensis, 223, 224, 337 cubanica, 48, 224, 337 juberthiei, 14, 224, 337 (Parvimysis) alrnyra, 11 (Parvimysis) bahamemis, 1 1 peckorum, 14, 223, 337 reddelli, 14, 337 (Surinamysis) americana, 11 Apogon rnenesemus, 248 novae-guineae, 248 savayensis, 248 snyderi, 248 Arachnomysis, 22, 28, 84, 327, 345 leuckartii, 121, 226, 327

641

TAXONOMIC INDEX

megalops, 75, 174, 233,. 327 Archaeomysis, 20, 41, 131, 220, 325, 345, 633 grebnitzkii, 42, 43, 146-149, 246, 249, 325 kokuboi, 325, 629 maculata, 157 Archoplites interruptus, 247 Arg yropelecus aculeatus, 531 hemigymnua, 531 Argyrozona argyrozona, 533 Aristeomorpha foliacea, 250, 534 Aristeus antennatus, 534 Arks felis, 244, 630 Amoglossws laterna, 241 Arripis trutta, 532 Artemia, 95, 261, 262, 447, 448, 457, 458, 465, 489 Arthromysis, 33, 337, 345 magellanica, 337 Arthrophryxus beringanus, 256 Ascarophis sp., 541 Asconiscidae, 257 A sconisccus simplex, 256 Aspidoecia normani, 255 Aspidophryxus, 252 frontalis, 256 peltatus, 256 Aspitrigla cuculus, 245 obscura, 245 Astroboa nigra, 186 Atheresthes stornias, 530 Atlanterythrops, 26, 327, 345 crassipes,. 327 Australerythrops, 26, 327, 345 paradicei, 327 Australomyeis, 31, 249, 332, 345

acuta, 332 incisa, 332

B Bairdiella chrysura, 243, 244, 631 Bartholomea annulata, 185, 186 Bathygadm melanobranchus, 630 Bathymysis, 30, 31, 88, 128, 332, 345 helgae, 332 renoculata, 184, 332 Bembras japonicus, 247 Bentheupha&, 384, 597 amblyops, 384, 429-431, 444, 445, 448,450, 457, 478,479,496, 513, 529, 530, 632, 636, 597 Bentheuphausiidae, 384 Benthosema glaciale, 531 suborbitale, 53 1 Biddulphia catenata, 458 Bolbosoma menoforme, 541 Bolinichthys longipes, 6 10 Boops boops, 241 Boreomysinae, 19, 20, 39, 322 Boreomysis, 9, 12, 19, 90, 96, 115, 123, 147, 220, 226, 238, 239, 250, 251, 322, 345, 633 acuminata, 332 arctica, 48, 57, 62-64, 102, 104, 106, 114, 116, 131, 132, 190, 200, 207, 227, 229, 239, 240, 250, 256, 322, 628 atlantica, 322 bispinosa, 322 brucei, 174, 235, 322 caeca, 322 californieu, 75, 79, 99, 101, 102, 104, 115, 128, 146, 148, 149, 322 californica longirostris, 11 chelata, 322 curtirostris, 76, 84, 322 dubia, 322 fragilis, 322

642

TAXONOMIC INDEX

hameni, 322 ill@, 226, 322 incisa, 76, 84, 322 inermis, 128, 227, 235, 322 insolita, 226, 322 interrnedia, 322 jacobi, 322 kistnae, 322 latipes, 322 longispina, 322 macrophthalina, 322 megalops, 48, 245, 322, 630 microps, 75, 82, 83, 87, 88, 114, 229, 322 nobilis, 194, 200, 227, 228, 322 obtusata, 322 plebeja, 76, 84, 322 richardi, 12 rostrata, 9, 12, 75, 85, 174, 322 rostrata japonica, 11 rostrata orientalis, 11, 12 rostrata var. Illig, 11 scyphops, 129, 130 semicoeca, 76, 84, 322 sibogae, 84, 256, 322 sphaerops, 322 tanakai, 12, 322 tattersalli, 322 tridens, 184, 229, 239, 322 tridens lobata, 11 vanhoeffeni, 12, 322 verrucosa, 322 Bowaniella, 14, 20, 325, 346, 633 atlantica, 14, 325 bacescui, 48, 325 banneri, 325 brasiliensis, 48, 77, 157, 161, 165, 166, 194, 325 dissimilis, 172, 325 $oridana, 325 inarticulata, 14, 325 johnsoni, 48, 325 merjonesi, 325 mexicana, 325 parageia, 172, 325 portoricensis, 49, 184, 325 reevensis, 14, 325 sewelli, 49, 172, 325 Brachyistius frenatus, 248 Brmidae, 530

Branchiophryxus caulleryi, 540 koehleri, 540 nyctiphumze, 539, 540 Brasilomysis, 28, 332, 346 castroi, 49, 156, 157, 161, 165, 166, 172, 194, 332 inermis, 332 Bregmacerotidae, 531 Brevoortia tyrannus, 243 Brotula multibarbata, 248 Bulla, 250

C Caesaromysis, 23, 28, 128, 327, 346 hispida, 75, 174, 220, 226, 327 Calanus finmarchicus, 603 tonsus, 528 Callionymus lyra, 241 Calyptomma, 28, 123, 332, 346 puritani, 231, 332 Caranx hippos, 630 Cardisoma guunhumi, 186 Carnegieomysis, 33, 128, 337, 346 xenops, 337 Caspiomysis, 33, 337, 346, 633 knipowitschi, 232, 337 Cassiopea androrneda, 186 Cathuracta antarctica, 535 Centropogon australis, 249 Centropristes striata, 243 Ceratolepis, 19, 320, 346 hamata, 49, 77, 220, 320 Ceratomysis, 14, 19, 123, 321, 346 egregia, 321 ericula, 14, 321 spinosa, 321 Ceratoscopelua warmingi, 531, 610 Chaetoceros, 449

643

TAXONOMIC INDEX

Chaetodon corallicola, 248 milearis, 248 Chalaraspidum, 19, 85, 128, 320, 346 alatum, 76, 220, 250, 320 Chalinura leptolepis, 238 Champsocephalus gunnari, 6 11 Charadriua

alexandrinus, 249 hiaticula, 249 Chauliodidae, 531 Chauliodus dame, 531 Chlamydomonas, 458 Chlorophthalmus agassizi, 610 Chlorotocus crassicornis, 534 Chromis agiii~,248 atripectoralis, 248 meruleus, 248 ovalis, 248 punctipinnis, 533 uerater, 248 Chunomysis, 28, 327, 347 diadema, 327 Citharichthys sordidua, 246, 529, 610 Cladocera, 96 Clupea harengus harengus, 240, 529 Clytia johnstoni, 257 Coccolithus huxleyi, 458 cocius crocodilus, 247 Coelorhynchus coelorhynchus, 610, 630 Cololabis saira, 532 Contracaecum sp. (Type B), 541 contracaecum sp. (Type D), 541 Coryphaenoides carapinua, 611, 630 Jilqer, 238 rupestris, 610 Coscinodiscus, 449

granii, 458 cottus asper, 246 Crangon crangon, 250 franciscorum, 631 septemspinosa, 250 Crustacea, 96 Cubanomysis, 31, 88, 332, 347 jimenesi, 49, 172, 332 Cumacea, 4 Cyathocephalus truncatus, 255 Cyclorrhynchus psittacula, 249, 535 Cyclotella ~ n a458 , Cynoscion arenarius, 244 nebulosus, 244, 630 Cypridina, 481

D Dactylamblyops, 27, 88, 128, 327, 347 fervida, 327 goniops, 327 hodgsoni, 174, 235, 327 iii, 327 laticauda, 84, 327 latisquamosa, 327 murrayi, 327 pellucida, 128, 327 sarsi, 327 solivaga, 327 stenurus, 327 tenella, 128, 327 thaumatops, 327 Dactylerythrops, 28, 123, 327, 347 bidigitata, 327 chrotops, 327 dactylops, 327 dimorpha, 327 gracilura, 327 Dajiidae, 257 Dajus, 252 mysidis, 256 siriella, 256 Daphnia, 91 pulex, 95 Daption capensis, 535, 611

644

TAXONOMIC INDEX

Daacyllua retkulatus, 248 Decapoda, 4 Diadema antillarum, 185 Diamysis, 12,13,35,337,347,633 americana, 12 aasimilis, 12 bahirensis, 49, 223, 225, 231, 232, 257, 337 bahirenais mecnikowi, 11 frontieri, 49, 337 pengoi, 49, 223, 225, 232, 337 pusilla, 232, 337 Diaphus dumerili, 531 lucens, 610 taaningi, 531 theta, 610 Diapterus plumieri, 630 Diaptomus ashlandi, 626 Dicentrarchus labrax, 240 Dichelopandalus bonnieri, 250, 534 Dinoffagallata, 251 Dinophyceae, 539 Diogenichthys laternatus, 610 Diomedea bulleri, 61 1 chlororhynchus, 250 exulans, 535 melanophris, 535 Dioptromyeis, 30, 88,128,332,347 paucispinosa, 172, 332 perspicillata, 332 proxima, 332 spinosa, 172, 332 Diplodua cervinus, 245 sargua, 245 Distomum $&jerum, 541 Doxomysis, 16, 31, 88,332, 347, 633 anonaala, 332 australiensis, 332 hanaeni, 332 littoralis, 332

longiura, 332 microps, 332 q d r k p i n o s a , 332 valdiviae, 13 zimmeri. 332

E Echinomysides, 23, 84, 128, 328,347 typica, 328 Echinomysis, 23, 128, 328, 347 chuni, 226, 328 distinguenda, 328 Echinorhynchus corrugatus, 541 salmonis, 255 Ectocarpus, 251 Electrow antarctiea, 532, 610 Ello biocystis my&arum, 253, 254 Ellobiopsidae, 253 Engraulis encrasicolus, 246 Entelurus aequoreus, 240 Eoerythrops, 26, 88, 328, 347 amamiensis, 328 typicus, 328 Eopsetta jordani, 246 Epistylus, 253 Erythropini, 22, 24, 88, 326 Erythrops, 14, 25, 128, 239, 240, 245, 328, 348, 633 abyssorurn, 227, 229,255, 328 africana, 328 bidentata, 14, 328 elegans, 45, 49, 61, 74, 96, 161, 166, 166, 171, 176, 194, 229, 240,241, 255, 256, 328 erythrophthdma, 49, 62, 104, 114, 115, 184, 228, 229, 240, 241, 244, 245, 256, 328, 626 frontieri, 328 glacialis, 228, 328 m i c r o p , 229, 255, 256, 328 minuta, 328 nana, 328 neapolitana, 328 parva, 49, 172, 328 peterdohrni, 231, 328

TAXONOMIC INDEX

serrata, 45, 46, 49, 61, 96, 114, 128, 129, 136, 161, 166, 168, 171, 176, 194, 208, 214, 240, 241, 255, 256, 260, 328 yongei, 328 Eucarida, 443 Euchaetomera, 23, 128, 328, 348 glyphidophthalmica, 75, 226, 328 intermedia, 174, 226, 328 oculata, 328 plebeja, 234, 328 richardi, 231, 328 tenuis, 75, 104, 114, 174, 226, 328 typica, 75, 174, 226, 328 zurstrasseni, 174, 328 Euchaetomeropsis, 23, 128, 328, 348 merolepis, 75, 226, 328 paciifica, 328 Eucinostomus argenteus, 630 gula, 630 Eucopia, 6,12,17,84,85,90,99,104,106, 120,123,131,133,220,321,348 australis, 84, 121, 128, 169, 174, 226, 235,249,256,321 grimaldii, 49, 62, 103, 114, 128, 169, 174,226,229,250,321 hanseni, 7, 10, 39,45, 55, 77, 128, 215 limguicauda, 321 sculpticauda, 49, 62, 83, 86-89, 91, 101, 103, 105, 106, 108, 114, 115, 117,134-136,141,169,174,226, 32 1 unguiculata, 45, 55, 60, 75, 77, 114, 115, 117, 128, 175, 215,226,250, 321 Eucopiidae, 17, 18, 321 Euma,lacostraca,4 Euphausia, 379,391,420,421,423,426, 427,456,478,504,526,530-532, 538, 546, 596,597, 609, 611 americana, 379, 382, 391, 432, 439, 450, 469,471,481,482, 536, 596, 597 brevis, 379, 392, 423, 427, 432, 439, 450,458,469,470,481,482,496 517,518, 522, 523, 525, 536, 597 crystullorophias, 379, 394, 423, 427, 432,450,470-472,483,525,536,

596,597,599,600,605,608 diomedeae, 375, 379, 393, 422, 423,

645

427,432,446,450,454,456-458, 478,479,489,496,503,511,521-

523,525429,531, 536,540, 596 distinguenda, 379, 382,395-397,422, 426,433,450,536,596 eximia, 379, 382, 391, 432, 450, 496, 501,518,520,525,536,596 fallax, 379, 397-399, 423, 427, 433, 443,452,497, 503, 531,537, 596 frigida, 379, 394, 427, 429, 432, 450, 503, 525, 536, 540, 596, 597, 599, 600,608,609 gibba, 379, 397, 421, 433, 452, 497, 536,596 gibboides, 379, 397-399, 423, 427, 433, 443, 452, 469, 472, 478, 480-482,503,518,520,537,596,

598 hanseni, 379, 401, 423, 427, 433, 443, 452,537,596 hemigibba, 379, 382, 401, 421, 423, 427,433,436,452,478,481,482, 497,518,523,537-540,598,609,

610 japonica, 48 1 krohni, 374, 379, 392, 423, 427, 429, 432,438,446447,450,460-463, 468-471,479,485,486,496,501, 503,511,514,516-518,524,525,

529,532-534, 536,539,540,597, 609 lamelligera, 379, 397, 433, 450, 536, 596 longirostris, 379, 401, 427, 433, 452, 537,596 lucens, 379, 394, 427, 432, 450, 496, 503, 519, 520, 523, 525,528, 529, 533, 536,540,596, 597,605, 609 mucronata, 379, 382, 395, 433, 450, 465, 520, 528, 529, 532,536, 538, 596 mutica, 379, 392, 427, 432, 439, 450, 462,518,522,523,536,596 nana, 379, 381, 382, 394, 427, 432, 450,462, 497, 503, 521, 525, 527, 529,536,596 pacijica, 116, 379, 381, 382,394,421, 422,426, 427,432,436,438,440, 450, 458, 460-473, 476, 481, 485-488,491,493,497,498,500, 501,503,511,512,514,519-521,

646

TAXONOMIC INDEX

524, 525, 527,530, 532, 533, 536, 539, 540, 543, 545, 549, 551, 553, 696,597, 600-602,604, 606-608, 6 10-612 paragibba, 379, 399, 421, 433, 438, 452, 522, 523, 537, 596, 598, 605, 609 pseudogibba, 379, 399, 400, 421, 427, 433,452,522,537,540,596 recwva, 379, 393, 426, 427, 432, 450, 462,471,520,523,525-529,533, 536,540,596,605 sanzoi, 379, 397-399, 423, 427, 433, 452,503,522,537,540,596 sibogae, 379, 382, 395-397, 422, 423, 426, 427, 433, 450, 503, 521, 522, 536,540,596 sibogae (distinguenda),457, 525 similis, 375, 379, 395, 421, 427, 432, 450,462,463,469-471,481,519, 522, 523, 525, 527, 536, 539, 540, 596 similis var. armatu, 379, 395, 433, 450,536,596 spinifera, 379, 401, 427, 433, 452, 520,525,528,537,596 superba, 212, 373, 379, 382, 393, 427-429,432,444-450,456,459, 460-472,474-477,481,483,486, 487,491,492,495,496,499-511, 513,526526,528,529,532-536, 540,544,545,549-553,596,597, 599-613 tenera, 379, 394, 427, 432, 439, 450, 457, 481, 518, 523, 523, 528, 529, 531,536,538,540,596,598 triacantha, 375, 379, 401, 427, 429, 433, 438, 448,452,497, 503, 511, 534,537,596,599,600,605 vallentini, 375, 379, 394, 427, 432, 450,473, 503, 519, 520, 521, 525, 528,536,540,596-598 Euphausiacea, 3,373,374,383,491,492 Euphausiidae, 385 Euthynnus (Katsuwonus)pelamis, 238 Eutrigla gurnardus, 241,245

F Fragillariopsis, 449

Fratercula arctica naumanni, 249 Fucus, 184 Fulmaris glacialis glacialis, 249

G Gadiculus argenteus argenteus, 245, 529 Gadus morhua morhua, 240, 529 Gaidropsarus mediterraneus, 240 Gammarus, 259 pulex, 120 Gangemysis, 35, 337, 349 assirnilis, 13, 223, 261, 337 Gasterosteus aculeatus, 240, 246 Gastrosaccinae, 20, 21, 39, 324, 632 Gastrosaccus, 13, 20, 68, 88, 90, 132, 141, 153, 155, 158, 184, 185, 240, 249, 250, 325, 349, 391, 629, 632, 633 armatus, 71,72 australis, 325 bengalensis, 325 bispinosa, 624,632 brevifissura, 194,325 dakini, 156,161,165,166,194,325 dunckeri, 325 elegans, 325 erythraeus, 235, 325 formosensis, 325 gordonae, 325 hibii, 325 indicus, 325 kempi, 326 lobatus, 13, 45, 49, 71-73, 152, 194, 250, 254, 256, 326 lobatus var. armata, 11 longifissura, 624, 632 magnilobatus, 13, 231, 326 mediterraneus, 72, 141, 152, 231, 326 rnsangii, 49, 74, 155, 184, 326 muticus, 256, 326 normani, 13, 73, 161, 166, 171, 176, 194,231,246,256,326 ohshimai, 326 olivae, 326 pacificus, 326

647

TAXONOMIC INDEX

parvus, 326 pelagicus, 326 psammodytes, 45, 46, 49, 57, 61, 73, 95, 96, 153, 155, 157, 158, 182, 184, 185, 194, 250, 258, 326, 628 pusillus, 326 roscoffensis, 326 sanctus, 49, 74, 155, 177, 179, 180, 185, 194, 231, 240, 241, 249, 326 sanctus widhalmi, 11 simulans, 90-92, 95, 146-148, 256, 326 spinqer, 43, 45, 49, 61, 73, 79, 91, 92, 96, 104, 114, 133, 141, 152, 161, 165, 166, 171, 176, 184, 229, 231, 239-242,256,326 vulgaris, 7, 45, 46, 49, 57, 61, 64, 189, 194,210,212,213,250,326 Gempylidae, 530 Gennadaa valens, 534 Geryon longipes, 251 Gibberythrops, 26, 328, 349 acanthura, 328 brevisquamosa, 328 Gigantocypris muelleri, 250, 534 Glyptocephalus cynoglossus, 241, 610, 630 zachirus, 529 Gnathophausia, 3, 19, 57, 59, 60, 65, 68, 84, 85, 88, 99, 102, 104-106, 108-111, 117, 132, 147-149, 191, 207,320,350 aflnis, 320 elegans, 320 gigas, 49, 62, 75, 76, 84, 99, 101, 103, 105, 115, 146, 148, 169, 175, 191, 215,220,226, 229,238, 260, 254, 320 gracilis, 49, 62, 75, 99, 101, 103, 115, 146, 148, 192, 215,220,226,254, 320 ingem, 6, 7, 50, 62-64, 75, 76, 98, 99, 101,103, 115, 117-120,132,133, 142-149, 156, 157,175,191,193, 194,197,199,207,210,211-213, 215, 218, 220, 226, 238, 250, 254, 320 longispina, 50, 62, 121, 123, 128, 320

scapularis, 234,320 zoea, 50, 62, 76, 77, 81, 83, 85, 103, 114, 115, 119,121-123,146,156, 191, 192, 215, 226, 229, 250, 251, 254,320,630 Gobius niger, 241 Gobiosoma robustum, 631 Gonostoma atlanticum, 531 Gonostomatidae, 531 Gymnerythrops, 28, 328, 350 anomala, 328 macrops, 328 microps, 235, 329 Gymnodinium kowalewskii, 95 Gymnoscopelus nicholsi, 532 Gymnothorax moringa, 253

H Halobaena caerulea, 611, 631 Halosauropsis macrochir, 239, 530 Hansenomysis, 19, 88, 90, 123, 227, 321, 350 angusticauda, 235, 321 antarctica, 235, 32 1 armata, 321 birsteini, 234, 321 chini, 234, 321 falklandica, 175, 180, 321 fyllae, 321 lucqugus, 234, 321 menziesi, 234, 321 peruvknus, 234, 321 rostrata, 321 spenceri, 234, 321 tattersallae, 234, 321 tropicalis, 234, 321 violacea, 32 1 Haplostylus, 13, 14, 20, 326, 350 bacescui, 14, 231, 326 estaf ricana, 326 parerythraeus, 326 Harelzgula pemacolae, 630

648

TAXONOMIC INDEX

Hemimysis, 31, 88, 337, 350, 633 m a y a m , 173, 341 abyssicola, 116, 337, 628 microps, 89, 90, 341 anomala, 50, 223, 225, 232, 261, 337 minuta, 341 lamornae, 10, 50, 57, 61, 81, 90-94, nouveli, 341 104, 114, 121, 133,146-148,161, odontops, 186, 341 166, 168, 171, 176,229,231,232, pacifica, 341 337 panamaensis, 234, 341 lamornae mediterranea, 11 proxima, 341 lamornae pontica, 11 rubrocincta, 50, 341 serrata, 232, 337 siciliseta, 341 speluncola, 64, 144, 145, 207, singaporensis, 34 1 210-213, 224, 337, 628 tasmanica, 341 Hemisiriella, 20, 90, 323, 351 tattersalli, 341 abbreviata, 323 waitei, 341 gardineri, 323 xanbhops, 341 parva, 323 zeylanica, 50, 186, 341 pulchra, 323 Heteromysoides, 36, 128, 341, 351, 633 Hemitaurichthys cotti, 14, 224, 341 thompsoni, 248 spongicola, 341 Heteroerythrops, 25, 329, 351 Heterophryxus microps, 329 appendiculatus, 540 purpura, 234, 329 Hippoglossoides tunseii, 329 elassodon, 246 Heteromysini, 22, 37, 340, 632 platessoides, 241, 242, 529 Heteromysis, 14, 36, 39, 84, 88, 186, 340, Holmsiella, 26, 329, 351 351,633 afinis, 10, 11, 146, 329 actinbe, 185, 186, 340 anomah, 10, 149, 329 armoricana, 44, 50, 340 Holmesimysis, 624, 625, 632, 633 atlantidea, 340 Costata, 624, 632 bermudensis, 50, 186, 340 n d a , 624, 632 bermudensis cesari, 11 nudensis, 624, 632 bredini, 340 sculpta, 624, 632 brucei, 340 sculptoides, 624, 632 cotti, 14 Holocentrus digitata, 235, 340 diadema, 248 djspar, 340 xantherythrus, 248 disrupta, 340 Holophryxus eideri, 231, 340 alascensis, 256 elegans, 50, 173, 213, 340 Hoplostethus floridensis, 340 mediterraneus, 630 formosa, 50, 61, 73, 229, 240, 241, Huso 244, 250, 341 huso, 246 gerlachei, 186, 235, 341 Hymenocephalue gomezi, 186, 341 italicus, 610, 630 guitarti, 50, 341 Hyperamblyops, 27, 128, 139, 351 gymnura, 50, 186, 341 antarctica, 329 harpax, 50, 186, 341 japonica, 329 kossmanni, 234, 341 megalops, 329 lybiana, 231, 341 nana, 128, 329 macropsis, 341 Hypererythrops, 25, 128, 146, 148, 329, mariani, 50, 186, 341 351

649

TAXONOMIC INDEX

caribbaea, 329 elegantula, 329 richardi, 329 serriventer, 329 spinifera, 329 zimmeri, 329 Hyperiimysis, 31, 88, 332, 351 madagascariensis, 332 Hyperprosopon argenteum, 248 Hyporhamphus laticeps, 248

I Ichthyobdella borealis, 631 Ictalurus catus, 247 furcatus, 244 melaa, 247 nebulosus, 247 punctatus, 247 Idiomysis, 14, 31, 337, 351 inermis, 337 japonica, 14, 337 tsurnamali, 50, 186, 235, 337 Iiella, 20, 326, 352 kojiwensis, 326 Iimysis, 16, 31, 88, 332, 352 atlantica, 332 orientalis, 332 Illex illecebrosus, 61 1 Indomysis, 35, 337, 352 ’ annandalei, 337 Inusitatomysis, 22, 31, 337, 352 insolita, 337 Isopoda, 4

J Johnius belengerii, 249 hololepidotus, 245

K Kainomatomysis, 33, 128, 337, 352 foxi, 231, 337 schieckei, 235, 337 Katamysis; 33, 337, 352, 633 warpachowskyi, 50, 223, 226, 232, 337

Katerythrops, 15, 25, 115, 329, 352 oceanae, 75, 136, 175,220, 226, 329 resimora, 329 tattersalli, 329 triangulata, 15, 329 Katsuwonua pehmis, 530

1 Labroides dimidiatus, 248 Lagodon rhomboides, 244, 631 Laminaria, 184 Lampanyctus australis, 6 10 cuprarius, 238, 531 mmropterus, 610 niger, 53 1 nobilis, 238, 531 regalis, 610 ritteri, 610 steinbecki, 53 1 Larus heermanni, 61 1 novaehollandiae, 61 1 novaehollandiae scopulinus, 535 Lates calcarifer, 631 Lauderia borealis, 458 Leiostomus xanthurus, 243, 244 Lepidomysidae, 16, 17, 39, 341 Lepidophanes guentheri, 531 Lepidorhombus boscii, 245 w h i m o n i s , 241, 630 Lepomis cyanellus, 247 gulosus, 247 macrochirus, 247 Leptomysini, 22, 29, 32, 331 Leptomysis, 13, 30, 88, 128, 177, 178, 236,332,352 apiops, 50, 181, 332 apiops banyulensis, 11 australiensis, 33 2 burgii, 62, 224, 231, 332 capensis, 332

650

TAXONOMIC INDEX

gracilis, 45,46,50,61,72,

73,96, 104, 115, 141, 162, 166, 169, 171, 176, 194, 207, 208, 214, 217, 229, 24Ck242, 245, 250, 254, 260, 332 lingwura, 13, 46, 50, 61, 62, 73, 99, 113, 138, 144, 145, 157, 162, 165, 166, 171, 176, 178, 181, 194,207, 208, 210-213,232,241,257,332, 628 mediterranea, 51, 91, 92, 104, 114, 155,181,333,627 megalops, 254, 333 peresi, 51, 231, 333 sardica, 13, 333 sardica pontica, 11 xenops, 333 Leptomysini, 16 Licmophora, 257 dalmatica, 257 lyngbyei, 257

Ligia oceanica, 120 Limanda limanda, 241 Limnocalanus macrurus, 626 Limnomysb, 35, 337, 353, 633 benedeni, 51, 223, 225, 232, 261, 337 Lobianchia dojleini, 531 Loligo opalescens, 6 11 Longithorax, 23, 220, 329, 353 alicei, 329 capensis, 175, 226, 329 fuscus, 83, 84, 86-89, 91, 329 nouveli, 329 similerythrops, 329 Lophogaster, 19, 84-86, 101, 10P106, 108-110,185,220,320,353,633 aflwis, 51, 320 americanus, 320 challengeri, 175, 320 erythraeua, 320 hawaiensis, 320 intermedius, 121, 320 japonicus, 320 longirostris, 129, 131, 320 multispinosus, 320 pacificus, 320 rotundatus, 321

schmidti, 321 spinosus, 77, 103, 114, 321 subglaber, 321 typicus, 7, 51, 62, 73, 81, 83, 84, 86, 87, 91, 103, 114, 121, 125, 133, 226, 245, 250, 321, 601, 609, 626, 63 1 Lophogastrida, 6 , 17, 90, 103, 104, 137, 320 Lophogastridae, 17, 18, 320

Lophogobius cyprinoides, 631 Lucania parva, 630 Lumpenus lampretaeformis, 609 Lycomysis, 35, 338, 353 bispina, 338 platycawla, 338 spinieawla, 338 Lyopsetta exilis, 529

M Macrocystis pyrifera, 95, 141 Macronectes giganteus, 535 Macrourus berglax, 610, 630 Maurolicua muelleri, 531 Megalactis hemprichi, 186 Meganyctiphanes, 379, 390, 423, 427, 478, 504, 530, 538, 597

norvegica, 379, 380, 390, 420-423, 425-427, 431, 437, 438, 440, 444-450, 456-473, 475, 476, 478-482, 484-486, 488-496, 499-501,503, 505, 510,511,514, 516, 517, 524, 525, 528, 629, 531-534,536,539,540,545,553, 597,601,602,604-611,613

Melanogrammus aeglejinus, 240,529 Melosira sp., 95 Menidia beryllina, 244, 631 menidia, 243

TAXONOMIC INDEX

Menticirrhus litoralis, 244 Merlangus merlangus merlangus, 240 Merluccius bilinearis, 243 capensis, 532 gayi gayi, 532 rnerluccius, 245, 609, 631 productus, 610 Memcanthomysis, 35, 338, 353 pygmaea, 51, 338 Mesomysis, 13, 15 czerniavsky, 13 incerta, 13, 15 kowalewski, 13 Mesopodopsis, 35, 88,338, 353, 633 africana, 166, 194, 249, 338 africana madagascariensis, 11 qrientalis, 39, 43, 44, 51, 57, 61, 63-65,225,261,338 slabberi,45,51,68,74,77,91-93,96,99, 105, 106, 112, 113, 133, 162, 165, 166, 181, 184, 190, 194,225, 229, 231,232,240-242,250,258,338 zeylanica, 51, 338 Metamblyops, 26, 329, 354 macrops, 329 oculata, 329 philippinensis, 329 stephensoni, 329 Metamysidopsis, 15, 28, 86, 88, 333, 354,633 elongata, 42, 44,45, 51, 61, 64, 68, 73, 99, 101, 133, 137, 140, 143, 145, 146, 148, 151, 158, 162, 166,

651

robusta, 184, 220, 227, 228, 254, 329, 626 Michtheimysis, 118 Michthyops, 27, 123, 330, 354 parva, 239, 330 theeli, 227, 330 Microgadus tomcod, 243 Microgobius gulosus, 631 Micromesistius australis, 529, 610, 630 poutassou, 245, 529 Micropogon undulatus, 243, 244 Micropterus salmoides, 247 Molva byrkelange, 532 molva, 240 Morone interrupta, 244 saxatalis, 243, 247 Mullus barbatus, 245 Munida sarsi, 250, 534 perarmata, 251 Myctophidae, 531 Myctophum aurolaternatum, 610 brachygnuthos, 610 nitidulum, 6 10 spinosum, 610 Myoxocephalus octodecemspinosus, 242 177-179,189,190~194~206,207, scorpius scorpius, 241 213,216,218,230,234,333 Myripristis elongata atlantica, 11, 157, 162, 165, amaenus, 248 166,194,230 kuntee, 248 insularis, 51, 57, 156, 333 murdjan, 248 macaensis, 15, 333 pralinius, 248 mexicana, 333 violaceus, 248 munda, 244, 333 Mysida, 6, 17, 39, 104, 137, 321 paci#ca, 234, 333 Mysidacea, 3, 4, 512 swyti, 51, 55, 173, 333 Mysidae, lY, 24, 29, 32, 34, 37, 39, 322 Meterythrops, 25, 329, 354, 633 Mysideis, 30,250,333,354 japonica, 329 insignis, 333 megalops, 329 parva, 231, 245, 333 microphthalma, 75, 76, 84, 329 Mysidella, 38, 86, 90, 341, 354 PiCta, 83,84,86-88,114,175,226,329 americana, 341

652

TAXONOMIC INDEX

minuta, 51, 173, 341 m m ,341 tanakai, 341 typhlops, 341 typica, 341 typica mauritanica, 12 Mysidellinae, 22, 37, 39, 341 Mysidetes, 22, 28, 88, 333, 355 anomala, 333 antarctica, 235, 333 bruchylepis, 235, 333 crassa, 333 dimorpha, 175, 235, 333 farrani, 228, 333 hanseni, 235, 333 intermedia, 333 kerguelensis, 333 macrops, 175, 333 microps, 175, 235, 333 patagonica, 175, 333 peruana, 234, 333 posthon, 51, 175, 235, 333 posthon microphthalma, 11, 12 Mysidion, 255 abyssorum, 255 commune, 255 Mysidium, 36, 185, 338, 355 columbiae, 45, 51, 57, 61, 64, 140, 153, 162, 165, 166, 169, 173, 179, 180,194,208,338 gracile, 129, 138, 153, 173, 178, 185, 186,251,338 integrum, 45, 51, 173, 338 Mysidobdella borealis, 631 Mysidopsis, 30, 84, 86, 88, 236, 333, 355, 633 acuta, 173, 333 almyra, 45, 51, 173, 244, 261, 333, 626, 630 angusta, 45, 51, 61, 162, 166, 171, 176,333 ankeli, 51, 173, 333 arenosa, 51, 333 bahia, 45, 52, 262, 263, 333 bigelowi, 184, 194, 228, 333 bispinosa, 333 bispinulata, 52, 173, 333 brattstroemi, 173, 333 californica, 234, 334 camelina, 52, 334

coelhoi, 334 coralicola, 52, 184, 334 cultrata, 52, 173, 334 didelphys, 10, 52, 61, 74, 95, 96, 104, 106, 114, 115, 162, 166, 171, 176, 189, 194, 202, 207, 208, 245, 256, 260, 334 eclipes, 334 eremita, 52, 334 furca, 173, 334 gibbosa, 10, 45, 52, 61, 63, 74, 91, 92, 95, 96, 128, 129, 136, 162, 167, 171, 176, 178, 183, 184, 194,204, 208,214,217,229,258,334 hellvillensis, 334 indica, 334 intii, 334 japonicu, 334 kempi, 334 kenyana, 334 major, 334, 631 mathewsoni, 173, 334 mauchlinei, 52, 334 mortenseni, 52, 173, 334 mortenseni cubanica, 12 robusta, 173, 334 robustispina, 334 schultzei, 334 similis, 245, 334 suedafrikana, 334 surugae, 334 tuironana, 45, 52, 173, 334 tortonesei, 45, 52, 77, 156, 157, 162, 165,167,173,194,334 velijera, 45,52,173,334 virgulata, 52, 173,334 Mysimenzies, 28, 123, 330, 355 hadalis, 69, 234, 330 Mysinae, 20, 22, 39, 326 Mysini, 16, 22, 32-34, 37, 335 Mysis, 7, 36, 88,200, 240,241,250,338, 355, 633 amblyops, 232, 338 australe, 338 w p i a , 232, 338, 627, 628 gaspemis, 158, 176, 181, 207, 228, 338 Zitoralis, 43, 52, 129, 131, 153, 156, 156, 195, 197, 223,225,227, 229, 233,338.625.626.629 macrolepis, 232, 338

TAXONOMIC I N D E X

microphthalma, 100, 111, 232, 338 mixta, 9, 43, 45, 52, 61,'73, 74, 96, 100, 156, 163, 167, 168, 184, 195, 201,215,228,229,238-242,256, 338, 626 oculata, 9, 41-43, 61, 121, 129, 132, 155, 156, 195,197,227,229,233, 238, 240, 249, 256, 338 polaris, 227, 233, 256, 338 relicta, 9, 42, 43, 45, 52, 57, 60, 63, 64, 69, 73, 74, 76, 77, 79, 91, 95, 96, 98-102, 112, 117, 118, 121, 129, 131-133, 141, 142, 146-148, 150-156, 158, 165,185, 190, 191, 193, 195,197-199, 201, 207,213, 216,219,220,223,225,228-230, 232,237,240,253,255,260-262, 338, 626-628, 632 stenolepis, 45, 52, 57, 60, 61, 65, 96, 99, 102, 121, 183, 195, 201, 207, 216, 228, 236,242, 244, 263, 338, 625, 629, 631 (Auricomysis)stenolepis, 12

-

N Nanomysis, 36, 338, 356 insularis, 338 siamensis, 223, 338 Navicula , 25 7 Nematobrachion, 379, 405, 447, 456, 457,478,495,598 boGpis, 379, 408, 434, 438, 444, 445, 452, 454, 469,472,478,479,497, 516, 537, 598, 604 fexipes, 378, 379, 405, 434, 452, 478, 497,518,519,537 sexspinosum, 379, 408,409, 434, 441, 452, 457, 469-472, 482, 537, 598 Nematonurus armatus, 238, 532 Nematoscetis, 378, 379, 383, 403, 422, 424, 428, 447, 456,457, 478, 483, 494,502,515,521,530-532,538, 542,598 atlantica, 375, 378, 379,405,406,424, 428,434,452, 462,497, 518, 523, 537,598,604 dificilis, 375, 379, 381, 382, 403, 421, 424, 428, 433, 436,443,452,458, 462,478,479,495,497-500,503, 519, 520, 525, 528, 529, 532, 537,

653

539,540,598,609,610 gracilis, 377-379, 405, 407, 424, 428, 434, 438, 452,457, 521, 523, 531, 537 lobata, 379, 405, 406, 428, 434, 452, 537 megalops, 379, 382, 403, 424, 428, 434,444, 445,452, 461, 462, 467, 469, 479, 481, 482, 490, 494, 497-499,501,503,511,516-518, 523,525, 529, 532, 537,539, 540, 598, 604 microps, 378, 379, 382, 404, 421, 424, 428, 434, 438,452,457,462, 472, 480, 518, 531, 532, 537, 598, 604 tenella, 375, 378, 379, 381, 404, 424, 428,434,438,452, 454,482,497, 511,518,531,537,598 Nemichthyidae, 531 Neoheteromysis, 36, 341, 357 naiilleri, 231, 341 Neomysis, 35, 41, 63, 84, 98, 100, 106, 134-136, 236, 246, 338, 357, 633 americana, 7, 10, 45, 52, 60, 61, 64, 65, 73, 74, 77, 99, 104, 115, 131, 146-148, 152,154, 156,163, 167, 168, 177, 190, 195, 203, 219,228, 242, 243, 250, 260, 262, 338, 631 awatschensis, 42, 98, 99, 105-107, 109, 110,146150, 154, 156, 158, 220,223,246,247,338 czerniavskii, 85, 247, 338 ilyapai, 338 integer, 9, 10, 43-46, 53, 57, 61, 63, 64, 77, 91-96, 98-101, 103, 105-116, 121, 128,129, 132, 134, 146, 147, 149-151,153-155,157, 163, 165, 167, 169, 171, 176, 177, 181, 183, 195, 204, 208,214,217, 225,229,237,240-242,249,253, 258, 338 intermedia, 43, 45, 46, 53, 57, 61, 63, 95, 96, 99, 100, 101, 105-107, 113,116,154-156,191,195,207, 220, 223, 228, 247, 261, 262, 338 japonica, 39, 45, 53, 63, 64, 98, 100, 102, 156, 210, 212, 213, 261, 338 kadiakensis, 69, 236, 246,254, 338 mercedis, 43, 95, 96, 149, 154, 156, 158, 163, 167, 195, 220, 223,234, 24&248,338,631

654

'TAXONOMIC INDEX

mercedis (awatschensis), 45, 46, 53, 73,77

meridionalis, 338 mirabilis, 95-97, 102, 106, 107, 109, 110, 142, 147, 149, 178, 180, 183, 190,219,339,627 monticellii, 339 nukazawai, 105 orientalis, 339 patagona, 339 rayii, 82-84, 101, 150, 228, 233, 339, 627 sopayi, 339 spinosa, 105-107,109,110,339

Nephrops norvegicus, 250, 534 Nezumia aequalis, 610, 630 Nipponerythrops, 22, 23, 330, 358 typica, 330 Notolychnus valdiviae, 531 Notophryxus clypeatus, 256 globularis, 540 lateralis, 539 ovoid-, 256 Notothenia microlepidota, 630 rossi marmorata, 533, 611 Nototodarus sloani sloani 533 Nouvelia, 13, 88, 334,358 natalensis, 334 natalensis mombasae, 12 nigeriensis, 334 valdiviae, 334 Nybelinia surmenicola, 541 Nyctiphanes, 379, 390, 423, 427, 478, 494,504,538,597,611,613

australis, 379, 382,390,427,431,450, 469471,496,503,513,520,521, 524, 525, 530, 532, 533, 535, 536, 540,597,607 capemis, 379,390,427,431,450,483, 496, 523, 525, 526, 532, 536, 540,597,609 couchi, 374, 379, 390, 422, 423, 427, 431, 439, 450, 458, 469, 471, 481,494,496,499-501,503, 511,

512, 516, 517, 529, 533, 534, 536, 539,597,601 simplex, 379, 391, 427, 432, 438, 450, 496,501,503,520,528-530, 536, 538, 604

0 Oceanites oceunicus, 535 Ocyropsis crystallina, 535 maculata, 535 Oncorhynchus gorbuscha, 532 keta, 533 tshawytscha, 247 Ophryodendron, 253 Oplophorus, 481 Opsunus beta, 631 Osmerus eperlanus, 240 mordax, 242

P Pachygrapsus crassipes, 500 Pachymetopon blochii, 533 Pachyptil a desolata, 611, 631 turtur, 535 Pagophilus groenlandicus, 238,529 Palaemon adspersus var.fabricii, 250 macrodactylus, 631 serratus, 2 12 Palaeoacanthocephalan, 541 Pandalus borealis, 603 Paracanthomysis, 35, 339,358 hispida, 147, 151,339 kurilensis, 339 Paracineta, 253 Paralabrax clathratus, 248 Paralepididae, 530 Paralepis atlantica prionosa, 532

TAXONOMIC INDEX

Paralichthys dentatus, 243 lethostigma, 243, 244 Paralophogaster, 6, 19,85,321, 358 atlanticus, 321 glaber, 321 indicus, 32 1 intermedius, 235,321 macrops, 235,321 microps, 235, 321 sanzoi, 235,321 Paramblyops, 27, 28, 123, 128, 330, 358 bidigitata, 330 brevirostris, 235, 330 globorostris, 330 rostrata, 250,330 Paramysis, 13, 15, 33,90, 132, 133,228, 237,246,258,339,358,633 agigensis, 232,339 arenosa, 45, 46, 53, 61, 63, 96, 104, 113, 114, 163, 167, 171, 176, 195, 208,217,241,339 bacescoi, 53, 163, 167, 195,339 baeri, 190, 223, 225, 232, 246, 261, 339,629 baeri bispinosa, 12 bakuensis, 232,339,629 eurylepis, 232, 339 festae, 231, 339 grimmi, 15, 339 helleri, 231, 339 incerta, 13,232,339 inflata, 15,339 intermedia, 45, 46, 53, 73, 76, 141, 190, 191, 196, 219, 223, 225, 232,246,261,339,629 kessleri, 223, 225, 232, 339 kessleri sarsi, 12, 255 kosswigi, 232, 339 kowalewski, 190, 262 k r ~ y e r i53, , 196,232,339 Zacustris, 12, 223, 225, 230, 232, 261, 262,339,629 lacustris tanaitica, 12 lacustris turcia, 12 loxolepis, 100, 111,232, 339 nouveli, 43, 44, 53, 163, 167, 196, 339 pontica, 53,196,232,339 portzicensis, 339

665

proconnesia, 231,339 ullskyi, 45, 46, 53, 73, 76, 191, 196, 223, 225, 232, 261, 339, 629 Paronatrema, 541 P a r a n c h i d i m , 20,326,359 angusta, 326 Parapagurus pilosimanus, 250 Parapemeus longirostris, 250, 534 Parapseudomma, 27,123,330,359 calloplura, 227,245,251,256,330 Parastilomysis, 33, 339, 359 paradoxa, 247,339 Parathemisto gaudichaudi, 528,533 paeifica, 539 Parerythrops, 25,330,359 afinis, 330 bispinosa, 330 lobiancoi, 231, 330 obesa, 229,255,256,330 paucispinosa, 330 spectabilis, 227, 330 Parvilux ingens, 610 Parvim ysis, 12 almyra, 12 bahamensis, 12 Pasiphaea multidentata, 544 sivado,534 tarda, 250 Pelichnibothrium caudatum, 541 Pempheris schomburgki, 248 Pemeus aztecus, 261 Peracaridrt, 4, 443 Perm jlavescens, 244,246 jlwviatilis, 238 Percarina demidofi, 246 Petalophthalrnidae,'18, 19, 39, 137,321 Petalophthlmus, 19,123,322,359 armiger, 81, 82, 84, 85, 87-89, 91, 128,226,322 caribbeanus, 322 oculatus, 322

656 Phaeodactylum tricornuturn, 458 Philocheras echinulatus, 250,534 Phoca hispida, 238 Pholia guanellus, 24 1 Phycis blennoides, 246,529 chesteri, 239, 529 Platichthys pesusjlesus, 241 stellatus, 246, 247 Platymysis, 624, 632, 633 facalis, 624, 632 Plesionika heterocarpus, 250,534 rnartia, 534 Plesiopenaeus edwardsianus, 250,534 Pleurerythrops, 15, 25, 330,359 constricta, 15, 330 inscita, 330 secunda, 330 Pleurobrachia pileus, 250 Pleuromarnma, 449 Pleuronectes platessa, 24 1 Plotus alle, 249, 535 Podocotyle atomon, 255 Pollachius pollachius, 240 virens, 240, 244, 529 Polycheles typhbps, 250, 534 Pomacentrus pavo, 248 Pomatomus saltatrix, 243 Pomatoschistus rnicrops,241 minutus, 241 Pomoxis annularia, 247 nigromaculatus, 247 Poatoporeia afinis, 102

TAXONOMIC INDEX

Porichthys notatus, 481,610 Potamomysis, 12 assirnilis, 12 Pranesus insularum, 248 Praunus, 10, 33, 46, 63, 84, 95, 96, 106, 153, 177, 207, 216, 218, 228, 258, 339,395,633 flexuosus, 7-9, 39, 41-46, 53, 57, 60, 61, 63, 64, 68, 71, 79, 83, 54, 86-88, 91, 92, 94, 96, 99, 100, 105, 108, 111, 114, 115, 117, 119, 120, 121, 123.127-130,132-138, 140-143, 145,154,155, 157,164, 167, 169, 171, 176, 183, 188, 196, 202, 208, 209, 215-218, 229, 237, 240-242, 249, 253, 258, 261,262,339,627,628 inermis, 9, 42, 45-47, 53, 61, 63, 133, 142, 164, 167, 171, 176, 183, 189, 196, 208, 215, 217, 218, 229, 240-242,330,339 neglectus, 7-9, 45, 46, 53, 61, 63, 71, 96, 99, 131, 138, 157, 164, 167, 171, 176, 183, 184, 196, 208, 218,229,240,242,253,339 Priacanthus cruentatus, 248 Prionace glauca, 610 Prionornysis, 30, 334,360 aspera, 334 stenolepis, 334 Prionotus carolinus, 243 Prodajus gastrosacci, 256 lobiancoi, 266 ostendensis, 256 ovatus, 256 Processa canaliculata, 534 Promysis, 30,86,88,334,360 atlantica, 53, 157, 173, 184, 334 orientalis, 334 Proneomysia, 36,339,360 eriopedes, 339 fusca, 339 lingvura, 340 longipes, 340

657

TAXONOMIC INDEX

misakiensis, 340 ornata, 340 perminuta, 340 quadrispinosa, 340 sandoi, 340 mrugensis, 340 takitai, 340 tenuiculus, 340 toriumii, 340 wailesi, 340 Protista, 251 Protomyctophum tenisoni, 532 Psetta maxima, 241 Psettichthys melanostictus, 246 Pseudamblyops, 27, 128, 330,360 conicops, 330 Pseudanchialim, 20,326, 360 erythraea, 235,326,391 inermis, 326 pusilla, 236, 326 sibogae, 326 Pseuderythrops, 26, 330, 361 gracilis, 330 Pseudeuphausia, 379, 391, 427, 478, 494,504,538 latifrons, 375, 379, 382, 391, 427, 432,450,521,522,536,538 sinica, 379, 382, 391, 427, 432, 450, 496,503,536 Pseudochaenichthys georgianus, 611 Pseudochalaraspidum, 19, 85, 128, 320, 361 hanseni, 320 Pseudochirella polyspina, 534 Pseudomma, 22, 27, 88, 123, 245, 330, 361,633 afine,7,53,61,129,163,167,171,176, 180, 184, 196,229, 256, 330, 626 antarcticum, 235, 330 armatum, 175,235,330 australe, 330 belgicae, 235, 330 berkeleyi, 330 bispinicaudum, 330 brwisquamosum, 330 calmani, 175,330

chattoni, 231,245, 330 crassidentatum, 53,330 frigidum, 227, 330 intermedium, 331 izuensis, 331 japonicum, 331 kruppi, 331 lamellicaudum, 331 latiphthalmum, 331 longicaudum, 331 longisquamosum, 33 1 magellanensis, 331 marumoi, 33 1 matsuei, 331 minuturn, 331 multispina, 33 1 nanum, 331 okiyamai, 331 omoi, 33 1 roseum, 229, 239, 256,331 sarsii, 53, 175, 235, 331 schollaertensis, 53, 235, 331 surugae, 331 tanseii, 331 truncatum, 220,228,233,331 Pseudomysidetes, 15, 30, 334, 361 cochinensis, 13,334 russelli, 335 Pseudomysis, 30, 128,335, 361 abyssi, 335 dactylops, 335 Pseudopleuronectes americanus, 243 Pseudotolithus elongatus, 244 Pseudoxomysis, 31,88,335, 361 caudaensis, 335 Pterodroma macroptera, 535 Pteromysis, 26, 331,361 amemiyai, 331 Pufinus gravis, 611 griseus, 6 11 Pygoscelis papua, 249

R Raja (Raja)brachyura, 239,530 (Leucwaja)circularis, 239, 245

658

TAXONOMIC INDEX

(Raja)clavata, 239,245,530 erinmea, 243, 530 (Leucoraja)fullonica,239 georgiana, 533 (Raja)montagui, 239,530 (Leucoraja)naevus, 239 ocellata, 243, 530 (Dipturus)oxyrinchus, 245 radiata, 243,530 (Amblyraja)radiata, 239 senta, 243,530 Rhinonemus cimbrius, 240 Rhizosolenia, 449 Rhopelophthalminae, 19, 20, 39, 324 Rhopalophthalmus, 19, 324, 361, 633 africana, 244, 324 brisbanensw, 156, 164, 165, 167, 196, 324 chilkensis, 324 dakini, 324 egregius, 324 jlagellipes, 324 indicus, 324 kempi, 324 longicauda, 54, 244, 324 longipes, 324 macropsis, 324 mediterraneus, 231,324 orientalis, 324 tattersallae, 324 terranatalis, 324 Rissa tridactyla tridactyla, 349

s Salmo salar, 240,609 trutta trutta, 240 Sargassum serratifolium, 184 Scaphocalanus magnus, 534 Schistomysis, 33, 90, 135, 228, 236, 237, 258,340,361,633 assimilis, 231, 340 elegans, 232, 340, 629 kervillei, 45, 46, 54, 61, 77, 96, 164, 165, 167, 171, 176, 196, 208, 217, 340 ornata, 46, 54, 61, 74, 77, 79, 81, 82,

86, 89, 91, 92, 96, 104, 115, 132, 136, 164, 167, 168, 171, 176, 196, 202,207,208,229,240-242,340, 633 parkeri, 114, 340 spiritus, 45-47, 54, 57, 60, 61, 63, 73, 77, 91, 92, 96, 114, 136, 138, 157, 164, 165, 167, 169, 17.1, 176, 177, 196,204,206,208,217-219,229, 237,240-242,255,257,340 Schizopoda, 4 Sciaenops oceZlata, 630 Scolamblyops, 27, 123, 331, 362 japonicus, 331 oculospinum, 234,331 Scomber (Scomber)scombrus, 241 Scophthalmus q u o s u s , 243 rhombus, 241 Sebastes atrovirens, 248,533 mystinus, 248 serranoides, 248, 533 Sergestes atlanticus, 534 corniculum, 534 grandis, 534 henseni, 534 pectinatus, 534 sargassi, 534 splendens, 534 Seriphus politus, 248, 533 Siphonochalina siphonella, 186 Siriella, 15, 20, 86, 88, 90, 91, 132-134, 136,169,176,323,362,633 adriatica, 231,323 aequiremis, 99, 101, 256,323 aflnis, 323 anomala, 323 armata, 45, 54, 61, 91, 92, 94, 104, 114,139,157,164,167,169,171, 181,196,240,323 australis, 323 brevicaudatu, 184, 323 brevirostris, 323 castellabatensis, 15, 54, 323 chierchiae, 54, 73, 173, 323

TAXONOMIC INDEX

clausii, 7, 9, 54, 61, 72, 73, 104, 114, 141, 164, 167, 169, 171, 196,231, 232,240,254,257,323 conformalis, 323 dayi, 323 denticulata, 323 distinguenda, 323 dollfusi, 235, 323 dubia, 323 gibba, 323 gracilis, 323 halei, 323 hanseni, 323 inornata, 256, 323 jaltensis, 54, 61, 71-73, 91, 92, 114, 141, 152, 164, 167, 169, 171, 181, 231,232,240,323 jaltensis brooki, 12 jaltensis crassipes, 12 jaltensis gracilipes, 12, 253, 257 japonica, 323 japonica izuensis, 12 japonica sagamiensis, 12 jonesi, 86,323 lingvura, 323 longidactyla, 323 longipes, 184, 323 media, 323 melloi, 15, 323 mexicana, 323 nodosa, 323 norvegica, 54, 73, 104, 114, 128, 129, 164, 167, 171,245,256,323 okadai, 323 pacifica, 73, 95, 96, 141, 178, 184, 234,248,324 panamensis, 234,324 paulsoni, 324 plumicauda, 324 podoensis, 324 quadrispinosa, 324 quilonensis, 324 robusta, 324 roosevelti, 234, 324 serrata, 235, 324 sinensis, 324 singularis, 324 tudjourensis, 324 thompsonii, 54, 175,226,256,324 trispina, 324 vineenti. 324

659

vulgaris, 256, 324 vulgaris rostrata, 12 wadai, 324 watasei, 324 watasei koreana, 12 watasei macropsis, 12 wolfi, 324 Siriellinae, 19, 20, 39, 323 Solea lascaris, 24 1 Solenocera membranacea, 250,534 Somateria mollissima borealis, 249 Spelaeogriphacea, 4 Spelaeomysis, 16, 17, 88, 128, 341, 362, 633 bottazzii, 224, 341, 624, 629 cardisomae, 186, 223, 342 longipes, 43, 54, 91, 92, 94, 96, 121, 196,224,342 nuniezi, 224,342 olivae, 224, 342 quinterensis, 224, 342 servatus, 180, 224,342 Spinachia spinachia, 240 Spim'nchus thaleichthys, 246 Sprattus sprattus sprattus, 240 Squalus acanthias, 239 Stellijer Eanceolatus,243 Stenobrachius leucopsarus, 610 Stenotomus chrysops, 243 Sterna paradisea, 249 Sternoptychidae, 531 Sternoptyx diaphana, 532 Stilomysis, 36, 340, 363 camtschatica, 340 grandis, 228, 340 major, 254,340 Streptodajus equilibrans, 256 Stygiomysidae, 16, 17, 39, 342

660

TAXONOMIC INDEX

Stygiomysis, 16, 17, 84, 88, 128, 224, 342,363,633 holthuisi, 84, 85,224, 342 hydruntina, 224,342 major, 224,342 Stylocheiron, 379, 410, 424, 428, 430, 442, 447, 456, 478, 494, 502, 530-532,538,542,546,598 abbreviatum, 379, 417, 424, 428, 435, 438, 444, 445, 452, 469, 472, 476, 478, 480, 497, 503, 516, 517,522, 525,531,537,599 aflne, 375, 377, 379, 410-412, 428, 434, 452, 478, 479, 497, 518, 520-523,531,537,540,598 carinatum, 379, 410, 424, 428, 434, 439,452, 497, 503, 521, 523, 531, 537,540,598 elongatum, 379, 415, 428, 435, 444, 445,452, 476,479,497,618,520, 529-531,537,599 indicum, 379, 414,415, 435, 452, 521, 537 insulare, 379,413,414,435,452,537 longicorne, 375, 377, 379, 380, 415, 416,421,424, 428, 435, 444,445, 452, 479, 484, 497, 503, 511, 516-518, 521, 531, 537, 540, 599,604 m a x i m u m , 379, 417, 418, 424, 428, 435, 438, 444, 445, 452, 470, 476, 479, 497, 516, 517, 519, 530,531,537,604 microphthalma, 379, 413, 435, 452, 497,523,531,537 robustum, 379, 418, 419, 435, 452, 537 suhnzi, 374, 379, 413, 424, 428, 435, 452, 497, 603, 517, 518, 523, 531,537,598 Surinamysis, 12 Symbolophorus cal qorniensis, 6 10 Symphurus nigrescens, 246 Synaphobranchus kaupi, 530 Synerythrops, 26, 331,363 cruciata, 331 intermedia, 331 truncata, 331

Syngnathus am, 240 Systellaspis debilis, 534

T Taius tumi&ons, 532 Tanaidacea, 4 Taphromysis, 36, 340, 363,633 bowmani, 45, 54, 61, 98, 99, 102,223, 340, 625, 626, 629, 630 louisianae, 223,340 Tarletonbeania crenularis, 610 Tenagomysis, 16,31,234,335,363 chiltoni, 7, 225, 335 macropsis, 54, 335 natalensis, 13 nigeriensis, 13 novae-zealandiae, 237,335 producta, 335 robusta, 335 scotti, 335 similis, 13, 335 tanzaniana, 185,335 (Nouvelia)tanzaniana, 12 tenuipes, 335 thomsoni, 335 Terapon jarbua, 249 Teraterythrops, 26, 128, 331,363 parva, 331 robusta, 75,220,331 Tessarabrachion, 379, 401, 447, 478, 495,542 oculatum, 377,379,401,433,452,460, 462-464,478,479,501,519,537 Thalassoica anturctica , 535 Thalassomyces, 25 1-254 albatrossi, 254 bitowskii, 254 boschmai, 254 fagei, 539, 540 f mciatus ,254 marsupii, 251, 539 nouveli, 254 Thalassomysis, 23, 33 1, 363 sewelli , 33 1 tattersalli, 331

TAXONOMIC INDEX

Thaleichthys pacificus, 532 Theragra chalcogramma, 609,631 Thunnus, (Germo)alalunga, 238, 530 (Ne,othunnus)albacares, 238, 530 atlanticus, 238 Thyrsites atun, 530, 609,631 Thysanoessa, 379, 381, 401, 422, 423, 428, 429, 447, 457, 461, 468, 470, 478, 484, 504, 516, 527, 529, 531, 532,538,540,598,600,606 gregaria, 379, 381, 402, 421, 423, 428, 433, 436, 452, 467, 478, 479, 497, 499,511,511,520-526,528,537, 539, 540, 598 inermis, 375, 376,379, 381,402,420, 421,428, 433,438,440, 446, 447, 452,457,460-462,467-471,478, 486, 491,492,497, 500, 501, 503, 511, 513-517, 519,525, 527,529, 533,535,537, 539, 540,553, 598, 600,601,607-609. inermis f. neglecta, 375, 376,446,447 inspinata, 379, 381, 382, 402, 433, 453,501,525,533,537 longicaudata, 379, 381, 382, 402, 428, 433,444, 445, 452, 479, 490,491, 493, 497, 502, 503, 511, 512, 516-518, 525,529, 535,537, 539, 545,598 longipes, 377, 379, 381, 382, 402, 420, 423, 428, 433, 438, 452, 469-473,488,511,516,519,525, 527,532,533,535,537,540,598 macrura, 379, 381,403,428, 433, 452, 537, 538, 598-600, 608, 609, 611 parva, 379, 402, 433, 452, 518, 537, 598 raschi, 374, 379, 381, 383, 403, 421, 428, 429, 433,438, 440, 444, 445, 447, 452, 456, 457, 459-464, 467-472,476,418,479,481,482, 486, 487, 489, 492, 497, 499, 501,503,511,512,514-517,519, 525, 527, 529, 533, 535, 537, 539, 540, 545, 553, 598, 600-603, 607-609 spinifera, 379, 381, 401, 433, 452,

661

465, 478, 501, 511, 519, 520, 525, 527, 532, 533, 537, 610, 611 vicina, 379, 403, 428, 433, 452, 519, 529,537,538,598 Thysanopoda, 379, 382, 385, 423, 427, 429, 456, 478, 480, 495, 504, 513, 530-532,538,542,547,597 acut$rons, 379, 389, 425, 427, 431, 444, 445, 448,450, 479, 503, 511, 516,511,528-530,536,538,597 aequalis, 374, 375, 379, 382, 383, 386, 387, 421, 423, 427, 431, 441, 448,450,454,457,462,469-471, 496, 511, 517, 518, 522, 523, 531, 536,597 astylata, 374, 375, 379, 382-384, 386, 387,431,441,450,522,536 cornuta, 379, 389, 427, 431, 445,448, 450, 478-480, 491, 494-496, 511,513,536 cristata, 379, 385, 427, 431, 445, 448, 450,454,457,470,482,496,536 egregia, 379, 389, 427, 431, 445, 448, 450,478-480,491,494-496,511, 536 microphthalma, 379, 389, 427, 43.1, 450,469-471,530,536 monacantha, 379, 385, 423, 427, 431, 438, 441, 448,450, 454, 457,469, 471,478, 479,481,496, 511, 531, 536,597,604 obtusifrons, 379, 388, 431, 450, 469, 471,531,536 orientalis, 379-381, 388, 421, 427, 431, 438, 450, 454,496,536, 597 pectinata, 379, 388, 423, 427, 431, 438,441,450,454,457,496,536 spinicaudata, 379,389,427,431,445, 448, 450, 478, 479,491,494, 511, 513,536 subaequalis, 374,375, 382,386 tricuspidata, 379, 381, 385, 420, 422, 423, 427, 431, 443, 445, 448, 450, 454, 458,464,469, 471, 478, 479,494,496,511,514, 523,525, 531,536,597,604 Todarodes paci$cus, 533 Trachinus, vipera,241

662 Trachurus japonicus, 532 symmetricus murphyi, 610 trachurus, 241,609,631 Trachyrhynchus trachyrhynchus, 610,630 Trematomus borchgrevinki, 533 newnesi, 533 Trichiurus japonicus, 610 lepturus, 610 Trigla Iucerna, 241, 245 lyra, 245 Triglidae, 244 Trigloporus lastoviza, 245 Triphoturus mexicanus, 61 0 nigrescens, 531 Trisopterus esmarki, 240 Euscus, 240 minutus minutus, 240

TAXONOMIC INDEX

Troglomysis, 33, 128, 340,363, 633 vjetrenicensis, 224, 340

U Uria, 535 aelge, 61 1 grylle mandtii, 249 Urophycis $oridanus, 243 regius, 243 tenuis, 242

v Valenciennellus, tripunctulatus, 53 1 Vinciguerria nimbaria, 531

X Xenistius californiensis, 248

Z Zoothamnium, 253 Zostera, 182 marina, 184

Subject Index A Abdomen dorsal spine, 376 dorsal swelling, 378 length of, 191-193 preanal spine, 381 Acanthocephalan worm, 255, 541 Aden, Gulf of, 410, 415 Adenine nucleotides, 116 Adenosine triphosphate, 476 Adoption of larvae, 60-63 Adriatic Sea, 46, 62, 178, 182, 254, 517, 60 1 Affinity with other crustaceans, 3, 4 Africa, 221, 230 east, 234, 392 north-west, 215, 230, 393, 408, 518, 531,610,630 south, 14,46, 194,244, 245, 249, 250, 258, 390, 393, 415, 417, 523, 532, 533,609,624,631,634 west, 230, 249 Age at sexual maturity, 194-206, 510-513,608,629 Aggregation, 166-182, 437, 514, 524526, 535-538, 550-552, 601, 604, 611-613 bioluminescence of, 483 breeding, 166, 167, 614 composition of, 178, 181 densities in, 179, 613, 614 function of, 181, 182, 237 types of, 170,613-615 Agulhas current, 389,523 Alabama, USA, 223 Alaska, 227,249,535 Gulf of, 376 Albacore, 238,530 Albatross, 76, 250 black-browed, 535 wandering, 535 Aleutian Islands, 233, 238,249,511 Alewife, 243 Algae, 96, 258, 448-453, 458, 476, 489 epiphytic, 257

Alimentary tract, 60, 91-94, 447, 448 Aluminium, 100,463 American coast, 69, 184, 228, 230, 233, 376,388,530,608 Amino acid, 101, 112, 113, 133, 429, 472-475,477 Ammonia, 488 Amphipod, 97,259,262,450-453 Anatomy and histology, circulatory system, 117-121, 481, 484 egg, 599 excretory organ, 132,133 eye, 123,128-131,478-480,604 gill, 118, 120 gut, 91-94,447,448 muscular system, 121-127 nervous system, 117, 121, 123, 128, 130,131,134,138 photophore, 480-483 reproductive system, 39-43, 599, 607,608 Anchovy, 242,244, 246 Andaman Sea, 399,413 Androgenic gland, 43,420 Antarctic, 174, 175, 227, 235, 238, 249, 254, 369, 383, 504, 508-511, 523, 524, 528, 529, 532, 533, 538, 599-601,603,609,610 convergence, 524 Antennae, 44,133,134 scale (exopodite),7, 10,447, 629 second (antenna), 132, 133 Antimony, 100,466 Antiperistalsis, 94 Anus, 94 Aorta cephalica, 117-119,121 descendens, 120 Apostome ciliate, 253,539 Arabian Sea, 375, 385, 386, 389, 391, 392, 394, 399, 404, 405, 413, 415, 417,521 Aral Sea, 631,634 Arctic, 151, 196, 223, 227, 239, 365,

663

664

SUBJECT INDEX

Arctic-continued 377, 383, 403, 513, 515, 516, 519, 529,608 ringed seal, 238 Argentine, 519 Arsenic, 101,463,465,602 Arteria abdominalis, 117-1 21 Arteria lateralis, 118-120 Ash, 98,99,106,459-461 Assimilation efficiency, 95-97,468,488, 489,544 Astaxanthin, 114, 472, 475, 476, 480, 602 Atlantic, 226-231, 235, 238, 239 east, 13, 215, 228, 230, 231, 235, 241, 244, 249, 367, 393, 408, 517, 518, 531,610,634 north, 13, 228, 239, 365, 383, 516, 517,529,608,630,634 west, 13, 224, 228, 230, 235, 242-244, 366, 517-519, 530, 608, 609, 634 Auk, crested, 249,535 least, 249, 535 little, 249, 535 parakeet, 249,535 Australasia, 221, 226,233, 368 Australia, 156, 161, 164, 185, 194, 196, 234, 249, 391, 395, 397, 399, 469, 521,609 Avoidance of nets, 68 Azov Sea, 79,194,196,246

Bass-continued yellow,'244 Bathymetric distribution adult, 14, 48-54, 66-78, 220, 259, 430-442 juveniles, 77,78, 177, 178,479 larva, 428-429,437-438,600 Bathypelagic species, 48-58, 62, 63, 66, 68-72, 75, 76, 207, 212, 226, 250, 430-435,478,480,495,513 Bay of Bengal, 385, 386, 391, 392, 394, 399,401,404, 410,413,415, 521 Bay of Biscay, 10 Bay of Fundy, 229, 376, 611, 613 Beaufort Sea, 402 Bec ocellaire, 131 Behaviour adoption of larvae, 60-63 aggregation, 166-182, 237, 524-526, 535-538,550-552,611-615 burrowing, 184, 185, 629 endogenous rhythms, 72-74, 79,141, 158, 439 substrate preference, 178, 181, 184, 185 Bellonci, organ of, 130 Beluga, 246 Bengal, Bay of, 385, 386, 391, 392, 394, 399,401,404,410,413, 415, 521 Bering Sea, 233,376,535,609,631 Bering Straits, 402 Bermuda, 534 Bib, 240 Bioluminescence, 132 chemistry of, 481 B Bacteria, 458,477,489,547 control of, 481-482 Baja California, 417, 512 effects of drugs on, 480, 482 Baltic Sea, 79, 100, 155, 156, 161-163, experimental studies of, 481, 482, 165, 184, 195, 225, 229, 230, 232, 487 242,262,366,608,634 function of, 482,483 Barbados, 5 I 9 of swarm, 483,526 13arents Sea, 104, 239, 376, 390, 417, periodicity of, 482 438,529,535,608, 613 spectral characteristics of, 481, 482, 13arium, 100 604 k r r a c o u t a , 530 Biomass 13ass euphausiid, 259, 430, 519, 542, 548, black sea, 243, 244 549,612 kelp, 248 mysid, 259, 548,549,629 largemouth, 247 Biometry, 8, 9, 189-193, 490-494, 606, sea, 240 607,624,629 striped, 243, 247 Biotin, 475

665

SUBJECT INDEX

Birds, 76,249,250,535-537,611,631 Biscay, Bay of, 10 Black Sea, 79, 194, 196, 225, 229, 231, 232,246,367 Blacksmith, 533 Blenny, snake, 609 Blood, 63, 115, 117, 120, 121, 471, 627 circulation, 117-121, 481,484 osmoregulation of, 121, 627 Bluefish, 243 Bluegill, 247 Blue whale, 528 Body density, 141,47 1,604 fluid, 471 length, 6, 10, 14, 15, 45-58, 61-62, 189-193,422,490-494,496-498, 500, 513, 543, 544, 604, 607, 629 shape, 10,502 surface area, 487,492 volume, 491,607 weight, 98, 106, 148-150, 189-193, 459,490-494,500,543,544,607, 627 Bogue, 241 Bopyrid parasite, 255, 256, 539, 540 Boron, 463 Bosphorus, 231 Bothnian Bay, 229, 230 Bouvet Island, 609 Brassicaterol, 111 Brazil, 14, 230, 156, 161, 162, 194 Breeding, aggregation, 166-170, 524, 525 areas, 599, 600 behaviour, 74 season, 45, 46, 166, 187, 194-206, 511-513,607,608,629 Brill, 241 British Columbia, 532 British Isles, 141 Brood, adoption of larvae, 60-63 lamellae, see Oostegites pouch. see Marsupium size of, 45-58,218,470,490,494-499, 513,606,624,625 Bryde’s whale, 526, 528 Bullhead, black, 247 brown, 247 Burma, 394

Butterfish, 241

C Cadmium, 101, 263, 463, 466, 602 Caecum, mid-gut, 94 Caesium, 102, 466 Calcium, 100, 461, 463 storage of, 94 California, 46, 77, 95, 156, 158, 162, 163, 177, 184, 216, 466, 469, 512, 514,532,610,611,631 Baja, 417, 520 current, 259,429,519,520,524 Gulf of, 520 Calorific value, 102, 103, 468, 470, 603, 627 Calyptopis, 383,420, 422,425,443, 501, 502 food of, 429 size of, 423,424 Canada, 13, 121,227,261,469,532,609 Canary Islands, 224,518,534 Cannibalism, 65, 534 Ca-pantothenate, 475 Cape Blanc, 391 Cape Cod, 242, 608, 609 Cape Hatteras, 530 Cape Verde Islands, 390, 402, 518 Capelin, 609 Carapace, lateral denticle, 380 length of, 189-193, 490-494 Carbohydrate, 94,98,99, 103, 106,460, 46 1 Carbon, 101,461,462,602,627 assimilation, 468,488 dioxide production, 150, 628 Cardiac cha,mber of stomach, 91-93 Caribbean Sea, 169, 172, 173, 223, 230, 231,235,244,519,624 Carotenoid, 114, 115, 472, 475, 476, 480, 602 Carrion, 97,258 Caspian Sea, 15, 100, 225, 229, 230,232, 246,367,629,634 Casts, see Integument Catfish blue, 244 channel, 247 sea, 630 white, 247

666

SUBJECT INDEX

Caves, 14, 17, 180, 213,224, 629 Cellulose, 96, 263 Cephalopod mollusc, 533, 536, 537, 611 Cerebral ganglion, 119, 121, 123, 128, 129,131 Cerium, 466, 547 Cestode worm, 255,541 Ceylon, 399 Chaetognath, 450-453,457 Channel Islands, 261 Char Lake, 151,195,197 Chemical composition, 98-1 16,459-477 amino acid, 101, 112, 113, 133, 429, 472-475,477 ash,98,99,106,459-461 astaxanthin, 114, 472, 475, 476, 480, 602 carbohydrate, 94, 98, 99, 103, 106, 460,461 chitin, 98, 99, 106,460,461,602 chitosan, 602 lipid (fat, oil), 94, 98, 99, 103-112, 459-461,468-472,603 metals, 100-102,461-467,602 other vitamins, 113-115, 472, 475 protein, 98-99, 106, 459-461, 472-475,603 vitamin A, 114, 115,472,476 water, 98, 99,459-461, 603 Chemical taxonomy, 9 Chile, 393, 520, 529, 530, 532, 538, 630 China, 223,260 Chitin, 98,99, 106, 460,461, 602 Chitosan, 602 Chlorophyll-a, 457 maximum layers, 441 Cholesterol, 111,472 Chromatophore, 16,115,132,183 Chromium, 100,463,464 Chromosomes, 43 Chukchi Sea, 233 Ciliatine, 603 Cladoceran, 95,97,632 Clyde, Firth of, 218,464,467, 512 Clyde Sea area, 10, 46, 161-164, 178, 438 Coastal species, 55-58, 61, 66, 68-71, 207,218,226-235,533 Cobalt, 463, 464 Coccolithophore, 448-453 Cod, 239,240,242,529,609

Coelenterate hydroid, 257 Colombia, 169, 230 Colour, see also Chromatophore change, 183 cryptic, 183 Columbia River, USA, 246 Commensalism, 185,186 Commercial exploitation, 260, 261, 549-551,553,611,612 Copepod, 95,97,447,457,534,603 choniostomatid, 252, 255 cyclopoid, 258 harpacticoid, 258 Copper, 100,463,464 Copper sweeper, 248 Coral Sea, 526,528 Costa Rica, 223 Crab-eater seal, 529 Crappie black, 247 white, 247 Crevalle jack, 630 Croaker, Atlantic, 243, 244 Crozet Island, 528 Cryptoxanthin, 476 Ctenophore, 250,535 Cuba, 14,224 Culture of mysids, 68,262

D Dab, long rough, 241 Dactyl, thoracic leg, 89, 90, 133, 139 Davis Straits, 517 DDT residue, 467,468, 603 Decapod crustacean, 250,533,534,536, 537,602 Deep-scattering layer, 69, 439, 440, 601,626 Delaware, USA, 77, 163, 195, 242, 250 Denmark, 7, 63, 95, 163, 164, 194, 196, 202,218,242 Density of adults, 141,471, 604 eggs, 60,429 population, 177, 179, 182, 260, 524, 526,542,613-615 water, 438 Desoxyribonucleic acid, 476 Detritus, 95,96,258,259,448-453 Development, 63-65,422,599,600

SUBJECT INDEX

Development+ontinued pathways of, 425 phases of, 59-64,207,208 Developmental ascent of larva, 77, 168, 428,429,442,600 Diatom, 76, 84, 95, 96, 446, 448-453, 458 epiphytic, 257 Diet, 262,448-453,457,458 Digestive system, 91-94, 119, 447, 448 diverticula, 40,41, 91 gland, 92-94, 119 Diglycerides, 106,470 Dinoflagellate, 95, 96, 251, 448-453, 539 Distribution (geographical), 14, 220235, 384-419, 515-526, 597-599, 629 bathymetric, adults, 14, 48-54, 66, 68-78,220,259,430-442,626 changes in, 76-79 distributional types, 48-54, 69-72 freshwater species, 223-225 juveniles, 77, 78, 177, 178,479 larvae, 428,429,437,438,600 latitudinal, 166, 167, 194-196 literature, 220, 221, 364-369, 38&419,597-599,634 migration, 168 patchiness, see Aggregation restricted, 170, 177, 178, 181, 182 substrate preference, 178, 181, 184, 185 Diurnal vertical migration, 71-76, 79, 152, 178, 430-442, 454, 456, 457, 488,546,601-602,636 energy expenditure on, 76, 102, 441, 488 extent of, 71,74-76,79,431-435 larvae, 428,429,437,438 Dniestr River, 225 Dorsal keel, spine, swelling, 376-378 Dragonet, 241 Drum, red, 630 Duck, eider, 249

E East China Sea, 233, 368, 521,532 East Indian Archipelago, 395 East Wind Zone, 510 Ecdysis, 44, 59, 65, 133, 142-146,

667

Ecdysis-continued 209-213, 421, 464-466, 484, 489, 500-509,543,544,547 Echinoderm, 450-453 Echo-sounder, 69,439,440, 601, 626 Egg, 39,42,45,437,599 density of, 60, 429 depth of laying, 429,599 development, 63-65, 422, 599-600, 625 fertilization of, 43, 45, 422 growth of, 42,499 laying of, 421,499,513,600 mass, 499 membrane, 60 number produced, 45-58, 470, 490, 494-499,513,606 variation in size of, 45-54, 422, 495, 600 Eilat, Gulf of, 417,522 Ejaculatory duct, 40-43 Electric field, 550 Electron transport system (ETS), 150, 605,606,628 Ellobiopsid, 251-254, 539, 540 Embryo, 39,4449,625 number produced, 45-58, 470, 490, 494-499, 513, 606, 624, 625 variation in size of, 45,46,48-54 Enderby Land, 469 Energy budgets, 151, 152,605 English Channel, 516 Environmental factors, biogenic structures, 181 density of water, 438 dissolved oxygen, 149, 156, 481, 487, 606 light, 71, 76, 79, 138, 152, 153, 178, 439,485,600,601,628 moonlight, 76,439 phytoplankton, 77,441,457 polarized light, 153 pollution, 10, 76, 262, 263 pressure, 138, 157,438,485,486 rain storm, 180 salinity, 63, 77, 79, 147, 149, 150, 154-156,438,485,523,628 solar eclipse, 439 substrate, 178, 181, 184, 185 temperature, 63, 76, 77, 79, 144-150, 153, 154, 438, 485,489, 513, 523,

668

SUBJECT INDEX

Environmental factors-continued 606 thermocline, 76, 154,438,439 tidal currents, 77, 141, 157, 177, 181 viscosity, 438 wave action, 77, 158 Enzyme, 84, 150, 477, 481, 488, 489, 603,604 Epipelagic species, 48-58, 61, 66, 68-72,207,226,238,470,513 Escape reaction, 182,605 Estonia, 262 Europe eastern, 223,225 northern, 225, 227 southern, 223 Excretion, 150, 151,488,544 Excretory organ, 132,133 Exoskelton, 461,464-467,477,547,602 Expeditions International Indian Ocean, 515 Experimental animals, 262, 263 External morphology, 6-13, 81-88, 375 Eyes, 534 anatomy and histology, 7, 119, 123, 128-132,478,604 bilobate (divided), 128, 376,478 chemical composition, 114, 115, 466 image formation, 604 nauplius, 131 photophore of, 478 pigment., 114, 129-131,480 size of, 478,479 spectral sensitivity, 131 variation in size of, 7, 11

F Faecal pellet, 92, 94, 97, 457, 465, 467, 468,547 Fairy prions, 535 Faroes, the, 390 Fat, see Lipid, Oil Father lasher, 241 F a t t y acid, 106-1 11,470-472 Fecundity, 45-58,470,490,499 Feeding, 90,484 appendage, 81-91,443-447 behaviour, 74,446,447 carnivorous, 91,95,445-448,627 current,, 447 filter, 90,91,445-448, 457,626

Feeding-continued experiments, 448,602 herbivorous, 84, 95 mechanism, 447 omnivorous, 445-448 rhythms, 441,454-457 Fertilization of egg, 43,45,422 Finback whale, 238 Fin whale, 527, 528 Firth of Clyde, 218,464,467,512 Fish, 236-249, 450-453, 481, 529-533, 536,537,609-611,629-631 farms, 261 Flavoxanthin, 476 Florida, USA, 223, 230, 630 Flounder, 237,241, 242,247 arrowtooth, 530 southern, 243,244 starry, 247 stony, 246 summer, 243 winter, 243 Fluoride, 602, 603 Folic acid, 475 Food, 76, 95-97, 133, 134, 186,448-453 assimilation of, 95-97, 488, 489, 544 basket, 90,446,447 chain, 258,545,546 digestion of, 94 diurnal variation in, 95 in laboratory, 95, 133, 134, 457, 458, 602,627 of larvae, 429 seasonal variation in, 95 selection, 627 France, 163,194-196,224,468 Freshwater species, 207, 223-225, 237, 261,364,365, 634 Frontal heart, 118,119 organ, 131, 132 Fuertaventura, Canary Islands, 438, 514,518 Fulmar, 249 Fundy, Bay of, 229,376,611-613 Fungus, 539 Furcilia, 383,425-428, 501, 505,534 food of, 429

G Galactose, 103 Gallium, 100

669

SUBJECT INDEX

Gascogne, Gulfof, 250,511, 512, 534 Gastropod mollusc, 250 Genetic variability, 382, 596 Genital aperture, 39 Geographical distribution, 8ee Distribution Geographical forms, 7,10,375-378 Georges Bank, 74,156,177,195,260 Gill, 118, 120 Gland androgenic, 43,420 digestive, 92-94, 119 endocrine, 132,420 excretory, 132,133 integumental, 134-137,484 Glucose, 103 Glycogen, 94,103,481 Goby black, 241 clown, 631 code, 631 common, 241 crested, 63 1 Golgi complex, 43 Gonad, 39-43, 188, 420, 421, 494-498, 543,544,606 metabolic demand of, 544 Gonostomid fish, 531 Grand Banks, Newfoundland, 242, 529 Gravid female, 438 Gravity receptor, 59, 65, 137-139, 193, 263 Great Barrier Reef, Australia, 185 Great Lakes, North America, 69, 75, 77, 100, 102, 195, 197, 198, 202, 225,255,260 Great Slave Lake, 195,197 Greenland, 154, 194, 225, 227, 376, 513 Growth rates, 207-213, 490, 500-514, 543445,553,606,607,629 egg, 499 energy budget of, 151,152 larva, 500-504 ovary, 494,543,544 Guillemot, 535 Mandt's, 249 Guinea basin, 388 Gulf of, 518 Gulf of Aden, 410,415 Gulf of Alaska, 376

Gulf of California, 520 Gulf of Eilat, 417,522 Gulf of Gascogne, 250,511,512,534 Gulf of Guinea, 5 18 Gulf of Maine, 201, 630 GulfofMarseille, 161, 162, 164, 165 Gulf of Mexico, 230, 244, 383-385, 388, 389,391,394,408,417,419,519 Gulf of Morbihan, 422 Gulf of Oman, 391,408,410,415,522 Gulf of St Lawrence, 242,376,467,469, 516,517,529 Gulf of Riga, 230 Gulf of Taganrog, 246 Gulf of Thailand, 521 Gulf Stream, 299,518,608 Gurnard grey, 241,245 red, 245 rock, 245 shining, 245 yellow, 241, 245 Gut, 60,91-94,447,448 Gwangura, 244

H Haddock, 240,529 Haemocyanin, 115,147 Haemolymph, 115 Hake, 245,609,631 longfin, 239,529 silver, 243 southern, 243 spotted, 243 white, 236,242 Harp seal, 238,529 Harvesting euphausiids, 459, 549-553,611-615 Hatchet fish, 532 Hawaii, 234, 248,259 Heart, 40,41, 60, 117, 118,485 Heat conductivity, 603 Hepatic artery, 118, 119 Hepatopancreas, 467 Herring, 236,240, 242,529,609 Histology, see Anatomy Homarine, 116,476 Hook-nose, 241 Hormone, 132-1 33 Hottentot, 533 Humpback whale, 628, 538

472,

670

SUBJECT INDEX

Hunting of prey, 448 Hydrocholesterol, 111 Hydrogen, 101,463 Hypogean environment, 14, 17, 180, 213,224,629

I Icelttnd, 239, 630 Identification and key, 16-38, 382-383, 625 India, 15, 196, 223-225, 234, 261, 391, 415,631 Indian Ocean, 226, 234, 235, 249, 368, 369,383,521-523 Insect, 97 Integument, 461, 464-467, 477, 547, 602 Intersexes, 10 Intestine, 40,41 Invalid species, 12, 13, 382 Invwtebrates, predation by, 250, 251, 533-537,611,631 Ireland, 195, 390 Irish Sea, 194, 196, 390, 529 Iron, 100,463,464 Isle of Man, 162, 164,165 Isoenzyme polymorphism, 9 Isolume, 439 Isopod, 250 epicaridean, 252, 255,256, 539, 540 Israel, 155, 177, 185,249 Italy, 15, 224 Ivory coast, 391,518

J Jamaica, 14, 162, 165, 194,223-224 Japan, 13-14, 46, 102, 105, 184, 194, 195, 223, 227, 233, 235, 247, 259, 261, 368, 393, 397, 399, 469, 524, 602,609,612,634 Inland Sea of, 184 Sea of, 149, 219, 233, 368, 376, 469, 521,533 J a l a , 415 Jenny, silver, 630 Jugoslavia, 224 Juveniles, 7, 17, 61, 62, 187, 188

K Kara Sea, 376 Kerguelen Island, 14,528

Key to genera, 13, 16-38,625 to species, 13, 382, 383 Killifish, rainwater, 630 Kingfish, Gulf, 244 Kittiwake, 249 Knysna Estuary, 245 Kob, 245 Korea, 254,260,521 Kootenay Lake, 261 Kuril-Kamchatka Trench, 3, 105, 227, 233,259 Kurile Islands, 223 current, 410,608

L Labium, 81.84-86 Laboratory maintenance of euphausiids, 457,458,517 mysids, 68, 262 Labrum, 81 Lacinia mobilis, 83, 84,443 Lake Huron, 195,198 Lake Michigan, 75,195,202,255,260 Lake Ontario, 77, 195, 198 Lake Piiiijiirvi, 154, 165, 195, 198, 214, 216,219 Lake Pontchartrain, 244 Lake St Lucia, 249 Lake Superior, 195, 197 Lake Tithoe, 261,632 Lake Washington, 76,246 Lamellae, see Oostegites Laminia ganglionaris, 129, 130,478 Lance, sand, 243,609,630 Laptev Sea, 195, 198 Largest mysid, 6 Larva, 40, 41, 57, 420-429, 437. 443, 599,600 adoption of, 60-63 behaviour of, 60 body length of, 61,62,65,423-424 development of, 57, 59, 60, 63-65, 426,502,625 developmental ascent of, 77, 168, 428,429,442,600 developmental pathways of, 425-428 distribution of, 428, 429 duration of development of, 63-65, 625 food of, 429

SUBJECT INDEX

Larva+ontiinued growth rates of, 501-504 identification of, 383 mortality of, 60 mouthparts of, 429 phases of development of, 59-64, 207,208 vertical distribution of, 428, 429, 437,438,600 see also Furcilia, Metanauplius, Nauplius, Pseudometanauplius Lateral denticle, 380 Lead, 100,463,467,547,602 Leech, 63 1 Leopard seal, 529 Lesser Antilles, 224 Life expectancy, 194-207, 490, 499, 510-513,552,553,608 Lightldark reaction, 482 Light, effects of, 71, 76, 79, 138, 152, 153, 178,439,485,600-601,628 Light lure, 68, 153,437 Lindane, 263 Ling, 240, 532 Lipid, 94,98,99,103-112,459-472,603 acid value, 106, 107 density of, 106, 107 fractions of, 106-1 12,470-472 iodine value, 106, 107 larval, 59, 60 neutralization value, 106,107 refractive index, 106, 107 saponification value, 106, 107 seasonal variation, 105, 106,470,472 Lith, 59,65, 137-139, 193,263 Lithuania, 230,262 Liver, 467 Lobster, 476 Loch Creran, 161 LochEtive, 74,78,79, 161, 163, 164 Loch Ewe, 44,47,161-164,218 Loch Fyne, Scotland, 114 Longevity, 194-206, 207, 490, 499, 510-513,522,553,608 Long Island Sound, 195,219,242,260 Louisiana, USA, 223,244 Luciferin, 481 Luminescent organ, see Photophore

M Mackerel, 241, 242

671

Mackerel-continued horse, 241, 609, 631 jack, 532, 610 Macrourid fish, 238, 532, 610,'611, 630 Madagascar, 223,234,391,399,528 Madeira, 103 Magellan, Straits of, 235 Magnesium, 100, 461,463 Maine, Gulf of, 201, 630 Maintenace in laboratory, 68, 262, 457, 458,517 Mandible, 81-85, 142, 443-446 of larva, 443 palp, 81, 82 Manganese, 100,464 Marseille, Gulf of, 161, 162, 164, 194 Marshall Islands, 248 Marsupium, 39-44, 59-65, 187, 188, 251,255 duration of development in, 63-65, 625 emergence from, 65,625 fluid of, 63 Maryland, USA, 244 Massachusetts, USA, 262 Mating, 43, 44, 251, 421, 483, 513, 514 Maxilla, 81, 86-88,443 exopodite of, 132 Maxillule, 81, 86, 443 Mediterranean Sea, 7, 10, 14, 101, 128, 141, 215, 227-232, 235, 245, 246, 254, 367, 383, 386, 417, 464-467, 480,511,517,529,609,631,634 Medulla externa, 129, 130, 132 Medulla interna, 129, 130, 132 Medulla terminalis, 129, 130, 132 Megrim, 241,630 Melanin, 95, 114,480 Menhaden, 243 Mercury, 100,101,463,466,467,602 Mesopelagic species, 48-58, 62, 66, 68-72,75,76,207,226,478,480 Messina, Straits of, 602 Metal content, 100-102,461-467,602 Metanauplius, 420,422,429 size of, 423,424,599 Meteor Seamount, 618 Mexico, 14, 223, 224,230 Mexico, Gulf of, 102,230,244,383-385, 388, 389, 391, 394, 408, 417, 419, 519

672

SUBJECT INDEX

Migration horizontal, 168 ontogenetic, 77,168,428,429,442,600 Minke whale, 528 Mississippi Sound, 244 Mojarra spotfin, 630 striped, 630 Mollusc, 97,450-453 cephalopod, 533,536,537,611 Moluccas, 415 Molybdenum, 463 Monaco, 104 Monoglycerides, 106,470 Moonlight, 76, 439 Morbihan, Gulf of, 422 Morocco, 533 Morphology, external, 6-13,81-88,375 Mortality, of adults, 215-218,251 larvae, 60,216-218 Moulting, see Ecdysis Mouth, 81, 94 Mouthpart, 81-88 of larva, 429 Mullet, striped, 245 Muscle, 465,466, 602 abdominal, 121-127 gut, 91-94 thoracic, 121-1 27 Myctophid fish, 238,531,532, 610

N Nauplius, 420,422,429 size of, 423,424, 599 Nauplius eye, 131 Nematode worm, 539 Neritic species, 71 Nervous system, 117, 121, 123, 128, 130,131,134,138 Netherlands, 195 Nets, 66-68, 168, 169, 175, 436, 437, 550,600,611,625 avoidance of, 68,426 New Brunswick, USA, 242 New England, USA, 262 Newfoundland Banks, 158, 176, 181, 456,517,529 New Guinea, 397 New Hampshire, USA, 262 Newman River, 230,262

New York, USA, 112 New Zealand, 234, 238, 393, 394, 513, 520, 521, 528, 530, 532, 535, 597, 611,613 Niacin, 475 Nickel, 100,463 Niobium, 466 Nitrogen, 101, 102, 461, 462, 488, 602, 627 North America, 69, 223 North Atlantic, 13, 228, 239, 365, 383, 516,517,529,608,630,634 North Carolina, USA, 244 North Pacific, 228, 233, 246, 367, 519, 631,634 NorthSea, 161,162, 164,165, 168, 229, 232,377,516 Norway coast of 116, 376, 377, 392, 468, 511, 531 fjords of, 376, 377,438,476,514, 601, 609 Norway pout, 240 Norwegian deep, 516 Sea, 608 Notacanthid fish, 239 Nova Scotia, 229,530,608 Novaya Zemlya, 249,535

0 Oceania, 221, 233, 234 Oceanic species, 174-176,226235 Oesophagus, 91-93 Oil, see Lipid Okhotsk, Sea of, 233,247,377,438,519 Oman, Gulf of, 391, 408, 410, 415, 522 Ommatidium, 7, 128-130, 450-453,480 Ommochrome, 114,480 Ontogenetic migration, 77, 168, 428, 429,442,600 Oostegite, 39,44,60,62 Optic artery, 118, 119, 130 Opticnerve, 121, 130,478 Oregon, USA, 75,79,246,512,530,532 Osmoregulation, 121, 627 Ostia, 117, 120 Ostracod, 97,250,481,534 Ovary, 39-42,420,421,495-498,606 germinal site, 39 metabolic demand of, 544

SUBJECT INDEX

Ovary--continued rate of growth, 494,543,544 volume of, 495,498,606 Oviduct, 39-42,45 Oxygen consumption, 146-150, 484488,543,544,605

673

Phaeophytin, 457 Philippines, the, 233,405,415 Phosphatidyl choline, 472 Phosphatidyl ethanolamine, 472 Phospholipid, 106, 108,470,472 Phosphorus, 101,461,463,488 Photophore, 132,378,476,478,480 P enlarged, 378,483 Pacific, 226-228,233,234 histochemistry of, 481 central, 234, 247,520 histology of, 480,481 eastern, 233, 238, 247, 368, 519, 520, larval, 425,480 624,634 Phytoplankton, 77, 96, 441, 457, 458, equatorial, 227 488,626 northern, 228, 233, 246, 367, 519, Pigment 631,634 blood, 115, 147 north-western, 233 chromatophore, 115,183 southern, 249 eye, 129,130,131,480 western, 233, 246, 368, 383, 520, 521, photophore, 481 634 plant, 457 Panama, 230 visual, 131 Paragnath, 81,84-86 Pinfish, 244, 63 1 Parasite, 251-257,539-541 Pipefish Pars incisiva, 443-446 great, 240 Pars molaris, 83, 84,443-446 snake, 240 Passamaquoddy Bay, USA, 195, 201, Piper, 245 203,242,262,494,514,529,631 Plaice, 236, 237, 241, 242, 529 Patagonia, 630 Pleopod, 44,447 Patchiness of distribution, see Aggrega- Plover, 249 tion Plutonium, 102,467,547 PCB, 468,547 Polarized light, 153 Penguin, 249,250 Pollack, 240, 244, 609, 631 Penis, 40, 41, 43, 188 Pollutants, 10, 65, 100-102, 262, 263, Pentose, 103 462-468,602,603 Perch Polonium, 467, 547 kelp, 248 Polychaete, 96 Sacramento, 247 Polychlorinated biphenols (PCBs), 463 silvcr, 243, 244, 631 Ponto-Caspian, region, 261 walleye surf, 248 Poor cod, 240 yellow, 244, 246 Population Peritrophic membrane, 92 analysis, 187-219, 508-514, 543-545, Persian Gulf, 375, 522 552,553,607,608,629 Peru, 223,438 density, 177, 179, 182, 260, 524, 526, trench, 69, 227,234 542,613-615 Pesticides, 463,603 production, 218,545 Petasma, 375,377,378,396 seasonal changes in size of, 160-182 Petrel, 631 Potassium, 100,463 Antarctic, 535,611 Prawn, culture of, 261 giant, 535 Preanal spine, 381 grey-faced, 535 Predator, 218, 23C251, 527-539, Pintado, 535 609-611,629-631 Wilson’s storm, 535 baleen whales, 238, 527-529,536,537

674

SUBJECT INDEX

Predator+ontinued birds, 249,250,535-537,611,631 escape from, 251 fish, 236-249, 529-533, 536, 537, 609-611,629-631 invertebrates, 250, 251, 533-537, 611,631 . seals, 238, 529, 536, 537 Pressure, 138, 157,438, 485,486 Protease, 477, 603 Protein, 98, 99, 106, 459-461, 472-475, 603 Provitamins D, 111,472 Pruth River, 225 Puerto Rico, 224 Puget Sound, 69 Puffin, Spitzbergen, 249 Pyloric chamber, 91-93

Q

Quark, 229 Queenfish, 248,533

Rhythms endogenous, 72-74, 79, 141, 158, 439 feeding, 441,454-457 oxygen consumption, 628 Ribbon fish, 610 Riboflavin, 475 Ribonucleic acid, 476 Ribose, 103 Riga, Gulf of, 230 Rockfish kelp, 248,533 olive, 236, 248, 533 Rockling four bearded, 240 shore, 240 Roscoff, 195, 196 Ross Sea, 469 Rostrum, 7,494 Rotifer, 96 Russia, 196 Ruthenium, 466

S

R Radioisotopes alpha, 467 industrial wastes, 102,464,466 naturally occurring, 467 radioactive fallout, 102, 464-466 Rain storm, 180 R a t tail, benthic, 532 Ray blonde, 239 cuckoo, 239 sandy, 239,245 shagreen, 239 spotted, 239 starry, 239 thornback, 239,245 Red Sea, 226, 235, 369, 375, 391, 399, 417,517,522 Regeneration, 10 Reproductive organ, 39-43, 188, 420, 421,496498,543,544,606-608 Respiration, 146-150, 484-488, 543, 544,605,627 Respiratory current, 140 Retinular cell, 129, 130 Retro-peristalsis, 94 Rhabdom, 128-130 Rhodium, 466

Sacramento-San Joaquin River Estuary, 46, 77, 156, 158, 163, 631 Saguenay Fjord, 629 Saithe, 240, 529 Salema, 248 Salinity, 63, 77, 79, 147, 149, 150, 154-156,438,485,523,628 Salmon, 240,468,609 chinook, 247 chum, 247,533 kokanee, 261 pink, 532 Salt balance, 121 Sand dab, 246,529 Sand lance, 243,609,630 Sardine, 248, 630 Sargasso Sea, 518,608 Saury, 532 Scaldfish, 241,245 School, 170,613,615 Scotia Sea, 466,529,609 Scotland, 7, 10, 46, 63, 77, 78, 105, 155, 160-169, 171, 176-178, 184, 188, 194-196, 202, 214, 216, 218, 225, 260,390,516 Sculpin longhorn, 242 prickly, 246

SUBJECT INDEX

675

Scup, 243,244 Smelt-continued Sea of Azov, 79, 194,196,246' longfh, 246 Seal, 238, 529, 536, 537 Snoek, 609,631 Sea of Japan, 149, 233, 368, 376, 469, Sodium, 100,463 521,533 Solar eclipse, 439 Sea of Okhotsk, 233, 247, 377,438, 519 Sole Searobin, 243,244 flathead, 236,246 Secondary sexual character, see lined, 631 Petasma, Thelycum petrale, 246 in mysids, 9, 10 rex, 529 Sei whale, 527, 528 sand, 241,246 Selenium, 463,465, 602 slender, 529 Selwyn River, 238 Sonic scattering layer, 69, 439, 440, Seminal vesicle, 40-42 601,626 Sensilla, 133-137, 380, 381, 484 Sound, reflection of, 440,601 Sensory organ, 130 South Africa, 14, 46, 194, 244, 245, 249, pore x-organ, 130, 132 250, 258, 390, 393, 415, 417, 523, seta, 133-137 532,533,609,624,631,634 Sex ratio, 213-216, 513,514 South America, 221, 230, 244, 519, 520 Sexual dimorphism, 16, 39-44, 188, South China Sea, 233, 368, 399, 408, 375,378,480,482,483 415,419,521 Sexual maturity, age at, 194-206, South Georgia, 476,528,611,631 510-513,608 Spawning, 43, 45, 421, 422, 429, 499, Shad, 242,247 513,599,600 Shearwater Species great, 611 introduction, 261, 632 Shetland Islands, 390 transplantation, 262 Shoal, 170,181,613 Specific heat, 603 Shrimp, 477,631 Spermathecum, 421 Siberia, 197 Spermatocytes, 42 Sierra Leone, 244 Spermatophore, 420-422,494,513 Silicon, 100 Spermatozoa, 40-44,421,494 Silver, 100 Spitzbergen, 249 Silverfish, 533 Sponge, 96,624 Silverside, 243 Spot, 243,244 tidewater, 244, 631 Sprat, 240 Sinking rates of Spur dog, 239 adults, 441, 604 Squid, 533,536,537, 611 cast integuments, 547 Star drum, 243 faecal pellets, 547 Statocyst, 59, 65,137-139,193,263 Sinus Sterol, 106, 111,470,472 gland, 130, 132 Stickleback system, 120 fifteen spined, 240 Skate, 530 three spined, 240,246 little, 243 St Lawrence, Gulf of, 242, 376, 467, long-nosed, 245 469,516,517,529 smooth, 243,244 Stomach, 40,41,91-94,447,448 thorny, 243,244 contents, 95-97, 448-453, 626 winter, 243 Stony Lake, 151,195,201 Skua, Antarctic, 535 Sturgeon, 255 Smelt, 240, 242, 247 green, 247

676

SUBJECT INDEX

Sturgeon-continued white, 247 Straits of Magellan, 235 Straits of Messina, 602 Striated body, photophore, 481 Strontium, 100,102,463,465 Sub-genera, 11, 12 Sub-species, 11, 12 Substrate preference, 178, 181, 184, 185 Succinic dehydrogenase, 116 Suctorian, 253 Sumatra, 413 Sunfish, 247 Surinam, 223 Swarm, see Aggregation Sweden, 198,261 Swimming, 140-142,605 speed of, 140,141,605

Tristan da Cunha, 250 Trout, 240,261,268 sand seatrout, 244 seatrout, 242 spotted seatrout, 244,630 Tuna, 530 blackfin, 238 skipjack, 238,530 slender, 530 striped, 530 yellowiin, 238, 530 Turbot, 241 Tyrrhenian Sea, 5 17

U . Uranium, 467 Urea, 488 Uropod, 7,189- 193,494,629

T Taiwan, 397 Tantalum, 100 Tasman Sea, 393,469,530 Telson, 7-10,629 Temperature, 63, 76, 77, 79, 144-154, 438,495,489,513,523,606 isotherm, 76 Tern, Arctic, 249 Testis, 42, 43, 420, 494 Texas, USA, 223,629 Thailand, 223 Gulf of, 521 Thelycum, 378-380,421,596 Thermocline, 76, 154,438,439 Thoracic leg, 39, 81, 88-91,447 cleaning mechanism, 89 elongated, 90,447 endopodite, 88, 89, 140 exopodite, 88, 89, 140 parts of, 20 Tidal currents, 77, 141, 157, 177, 181 Tin, 100 Tintinnid, 96,448-453,457 Titanium, 100,463 Toadfish, Gulf, 631 Tocopherol, 472 Tomcod, 243 Trace metals, 486 Trematode worm, 255,541 Triacylclycerols, triglycerides, 106, 108, 111,470-472

V Vanadium, 463 Variation, 6-13, 375, 624 body shape, 10 colour, 16, 115, 132, 183 egg, size of, 45, 46, 48-54, 422, 495, 600 genetic, 382, 596 geographical form, 7,10,375-378 larval development, 425-428 lateral denticle, 381 metanauplius, size of, 423, 424, 599 morphological, 6-13 photophore, 378 preanal spine, 381 Vas deferens, 40-43,420 Venezuela, 156 Vertical distribution adults, 14, 48-54, 66-78, 220, 259, 430-442 bathypelagic specios, 48-58, 62, 63, 66, 68-72, 75, 76, 226,250, 43& 435,478,480,495,513 juveniles, 77, 78, 177, 178,479 larvae, 428,429,437,438,600 eggs, 429 epipelagio species, 48-54,431-435 mesopelagic species, 48-54, 431-435 size groups, 75, 77, 168, 428, 429, 442,600

677

SUBJECT INDEX

Vertical migration, 71-79, 102, 430442,454,456,457,626 of larvae, 428,429,437,438 of size classes, 75,438 physical factors controlling, 76, 77, 79,437-439,600,601 significance of, 440-442 Virgin Islands, 249 Virginia, USA, 104 Viscosity, 438 Vision, 129-131 image formation, 604 Vitamin, 113-115,472,475 Vitamin A, 114,115,472,476 Vitamin B,,, 113,475 Vitamin BT,475 Vitamin C , 113 Vitamin E, 472 Volga River, 46, 77, 225

w Warmouth, 247 Washington, USA, 246 Water consumption, 94 content, 98,99,459-461, 603 Wave Action, 77,158 Wax esters, 106, 108, 470,471 Weakfish, 243 Weaver, lesser, 241 Weddell Drift, 508, 510 Weddell Sea, 394 Weddell seal, 238 Well, 17, 224

West Wind Drift, 394 Whale, 76, 238, 527-529 extent of predation by, 536-537 feeding methods, 527 food of, 238,526-529 Whitefish, mountain, 261 White Sea, 516 Whiting, 240 southern blue, 630 worm, 255 Windowpane, 243 Witch, 241 World list of Mysids, 6, 12, 13-16, 221, 320-342,632 worms acanthocephalan, 255,541 cestode, 255, 541 trematode, 255, 541

X X-organ, 130, 132

Y Yellow Sea, 532 Yolk, egg, 59,60 Y-organ, 133 Yttrium, 465 Yucatan, 224

Z Zanzibar, 224 Zeaxanthin, 476 Zinc, 100,463-465,547,602 Zirconium, 466

This Page Intentionally Left Blank

Cumulative Index of Titles Alimentary canal and digestion in teleosts, 13, 109 Antarctic benthos, 10, 1 Artificial propagation of marine fish, 2, 1 Aspects of stress in the tropical marine environment, 10, 217 Aspects of the biology of seaweeds of economic importance, 3, 105 Association of copepods with marine invertebrates, 16, 1 Behavioirr and physiology of herring and other clupeids, 1, 262 Biological response in the sea to climatic changes, 14, 1 Biology of ascidians, 9, 1 Biology of coral reefs, 1, 209 Biology of euphausiids, 7, 1 ; 18, 373 Biology of mysids, 18, 1 Biology of pelagic shrimps in the ocean, 12, 233 Biology of Pseudocalanus, 15, 1 Biology of wood-boring teredinid molluscs, 9, 336 Blood groups of marine animals, 2, 85 Breeding of the North Atlantic freshwater eels, 1, 137 Circadian periodicities in natural populations of marine phytoplankton, 12, 326 Diseases of marine fishes, 4, 1 Ecology and taxonomy of Halimeda: primary producer of coral reefs, 17, 1 Ecology of Intertidal Gastropods, 16, 11 1 Effects of heated effluents upon marine and estuarine organisms, 3, 6 3 Estuarine fish farming, 8, 119 Fish nutrition, 10. 383 Floatation mechanisms in modern and fossil cephalopods, 11, 197 General account of the fauna and flora of mangrove swamps and forests in the Indo-West Pacific region, 6 , 7 4 Gustatory system in fish, 13, 53 Habitat selection by aquatic invertebrates, 10, 271 Heavy metals, 15, 381 History of migratory salmon acclimatization experiments in parts of the Southern Hemisphere and the possible effects of oceanic currents and gyres upon their outcome, 17, 397 Influence of temperature on the maintenance of metabolic energy balance in marine invertebrates, 17, 329 Interactions of algal-invertebrate symbiosis, 11, 1 679

680

CUMULATIVE INDEX OF TITLES

Laboratory culture of marine holozooplankton and its contribution to studies of marine planktonic food webs, 16, 21 1 Learning by marine invertebrates, 3, 1 Management of fishery resources, 6, 1 Marine biology and human affairs, 15, 233 Marine molluscs as hosts for symbioses, 5, 1 Marine toxins and venomous and poisonous marine animals, 3, 256 Methods of sampling the benthos, 2, 171 Nutritional ecology of ctenophores, 15, 249 Parasites and fishes in a deep-sea environment, 11, 121 Particulate and organic matter in sea water, 8, 1 Petroleum hydrocarbons and related compounds, 15, 289 Photosensitivity of echinoids, 13, 1 Physiological mechanisms in the migration of marine and amphihaline fish, 13, 248

Physiology and ecology of marine bryozoans, 14, 285 Physiolohy of ascidians, 12, 2 Pigments of marine invertebrates, 16, 309 Plankton as a fact,or in the nitrogen and phosphorus cycles in the sea, 9, 102 Pollution studies with marine plankton: Present statue of some aspects of marine microbiology, 2, 133 Problems of oil pollution of the sea, 8, 215 Rearing of bivalve mollusks, 1, 1 Recent advances in research on the marine alga Acetabularia, 14, 123 Respiration and feeding in copepods, 11, 57 Review of the systematics and ecology of oceanic squids, 4, 93 Scatological studies of the bivalvia (Mollusca), 8, 307 Some aspects of the biology of the chaetognaths, 6, 271 Some aspects of neoplasia in marine animals, 12, 151 Some aspects of photoreception and vision in fishes, 1, 171 Speciation in living oysters, 13, 357 Study in erratic distribution: the occurrence of the medusa Conionemus in relation to the distribution of oysters, 14,251 Taurine in marine invertebrates, 9, 205 Upwelling and production of fish, 9, 255

Cumulative Index of Authors Allen, J. A., 9, 205 Ahmed, M., 13,357 Arakawa, K.Y., 8, 307 Balaknshnan Nair, N., 9, 336

Lurquin, P., 14, 123 McLaren, I. A., 15, 1 Macnae, W., 6,74 Marshall, S. M., 11, 57

Rlaxter, J. H. S., 1, 262

Mauchline, J., 7 , 1 ; 18, 1

Boney, A. D., 3, 105 Bonotto, S.,14, 123 Branch, G.M., 17, 329 Bruun, A. F., 1, 137 Campbell, J. I., 10, 271 Carroz, J. E., 6, 1 Cheng, T.C., 5, 1 Clarke, M. R., 4, 93 Corkett, C. J., 15, 1 Corner, E.D. S., 9, 102; 15,289 Cowey, C. B., 10,383 Cushing, D. H., 9, 255; 14, 1 Cushing, J. E., 2, 85 Davies, A. G., 9,102; 15,381 Davis, H.C., 1, 1 Dell, R . K., 10, 1 Denton, E.J., 11, 197 Dickson, R.R., 14, 1 Edwards, C., 14, 251 Evans, H. E., 13, 53 Fisher, L. R., 7, 1 Fontaine, M., 13, 248 Garrett, M.P., 9, 205 Ghirardelli, E., 6,271 Gilpin-Brown, J. B., 11, 197 Goodbody, I., 12, 2 Gotto, R. V., 16, 1 Gulland, J. A., 6, 1 Harris, R. P., 16, 211 Hickling, C. F., 8, 119 Hillis-Colinvaux, L., 17, 1 Holliday, F.G. T., 1, 262 Kapoor, B. G., 13, 53, 109 Kennedy, G. Y., 16, 309 Loosanoff, V. L., 1, 1

Mawdesley-Thomas, L. E.,12, 151 Mazza, A., 14, 123 Meadows, P.S., 10,271 Millar, R. H., 9, 1 Millott, N., 13, 1 Moore, H. B., 10,217 Naylor, E.,3, 63 Nelson-Smith, A.,8, 215 Newell, R. C., 17, 329 Nicol, J. A. C., 1, 171 Noble, E. R., 11, 121 Omori, M., 12,233 Paffenhofer, G-A., 16, 211 Pevzner, R. A., 13, 53 Reeve, M. R., 15,249 Riley, G. A., 8, 1 Russell, F. E., 3, 256 Russell, F. S., 15,233 Ryland, J. S., 14, 285 Saraswathy, M., 9, 336 Sargent, J. R., 10, 383 Scholes, R.B., 2, 133 Shelbourne, J. E., 2, 1 Shewan, J. M., 2, 133 Sindermann, C. J., 4, 1 Smit, H., 13, 109 Sournia, A.,12, 236 Stewart, L., 17, 397 Taylor, D. L., 11, 1 Underwood, A. J., 16, 111 Verighina, I. A., 13, 109 Walters, M. A., 15, 249 Wells, M.J., 3, 1 Yonge, C. M., 1, 209

681

This Page Intentionally Left Blank