Atlas of Mammalian Chromosomes

ATLAS OF

Mammalian Chromosomes

ATLAS OF

Mammalian Chromosomes Edited by

Stephen J. O’Brien Joan C. Menninger William G. Nash

A John Wiley & Sons, Inc., Publication

Copyright © 2006 by John Wiley & Sons, Inc., Hoboken, NJ. All rights reserved. Published simultaneously in Canada No part of this publication may be reproduced, stored in a retrieval system, or transmitted in any form or by any means, electronic, mechanical, photocopying, recording, scanning, or otherwise, except as permitted under Section 107 or 108 of the 1976 United States Copyright Act, without either the prior written permission of the Publisher, or authorization through payment of the appropriate percopy fee to the Copyright Clearance Center, Inc., 222 Rosewood Drive, Danvers, MA 01923, (978) 750-8400, fax (978) 750-4470, or on the web at www.copyright.com. Requests to the Publisher for permission should be addressed to the Permissions Department, John Wiley & Sons, Inc., 111 River Street, Hoboken, NJ 07030, (201) 748-6011, fax (201) 748-6008, or online at http://www.wiley.com/go/permission. Limit of Liability/Disclaimer of Warranty: While the publisher and author have used their best efforts in preparing this book, they make no representations or warranties with respect to the accuracy or completeness of the contents of this book and specifically disclaim any implied warranties of merchantability or fitness for a particular purpose. No warranty may be created or extended by sales representatives or written sales materials. The advice and strategies contained herein may not be suitable for your situation. You should consult with a professional where appropriate. Neither the publisher nor author shall be liable for any loss of profit or any other commercial damages, including but not limited to special, incidental, consequential, or other damages. For general information on our other products and services or for technical support, please contact our Customer Care Department within the United States at (800) 762-2974, outside the United States at (317) 572-3993 or fax (317) 572-4002. Wiley also publishes its books in a variety of electronic formats. Some content that appears in print may not be available in electronic formats. For more information about Wiley products, visit our web site at www.wiley.com. Library of Congress Cataloging-in-Publication data is available. ISBN-13 978-0-471-35015-6 ISBN-10 0-471-35015-X

Printed in the United States of America 10 9 8 7 6 5 4 3 2 1

CONTENTS Foreword xxix Preface

xxxi

Acknowledgments Contributors

xxxv

xxxvii

SPECIES

COMMON NAME

2N

SOURCE*

MONOTREMATA

1

Order Monotremata

1

Family Tachyglossidae Tachyglossus aculeatus Tachyglossus aculeatus Zaglossus bruijni Zaglossus bruijni Family Ornithorhynchidae Ornithorhynchus anatinus Ornithorhynchus anatinus

Short-beaked echidna/Short-nosed echidna Short-beaked echidna/Short-nosed echidna Long-beaked echidna/Long-nosed echidna Long-beaked echidna/Long-nosed echidna

64F 63M 64F 63M

Graves et al. (Unpublished) Graves et al. (Unpublished) Graves et al. (Unpublished) Graves et al. (Unpublished)

6 6 7 7

Platypus Platypus

52F 52M

Rens et al. (2004) Rens et al. (2004)

8 8

MARSUPIALIA

9

Order Didelphimorphia

9

Family Marmosidae Marmosops incanus Marmosa sp. Micoureus demerarae Thylamys elegans Metachirus nudicaudatus Monodelphis domestica Monodelphis domestica Ideogram Family Caluromyidae Caluromys lanatus Family Didelphidae Philander opossum Didelphis marsupialis Didelphis virginiana

South American mouse opossum Mouse opossum Mouse opossum Elegant fat-tailed opossum/Southern mouse opossum Brown four-eyed opossum Short-tailed opossum Short-tailed opossum

14 14 14 14

Svartman (1998) Aniskin (1994a) Svartman (1998) Shchipanov et al. (1996)

12 12 13 13

14 18 18

Aniskin et al. (1991) Graphodatsky et al. (Unpublished) Pathak et al. (1993)

14 15 16

Woolly opossum

14

Casartelli et al. (1986)

17

Black four-eyed opossum Common opossum Virginia opossum

22 22 22

Svartman (1998) Aniskin et al. (1991) CRES San Diego (Unpublished)

17 18 18

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

v

vi

CONTENTS

SPECIES

COMMON NAME

2N

SOURCE* 19

Order Paucituberculata

Family Caenolestidae Rhyncholestes raphanurus

Chilean “shrew” opossum

14

Gallardo and Patterson (1987)

20

Order Microbiotheria

Family Microbiotheriidae Dromiciops australis

Monito del Monte

14F/13M

Gallardo and Patterson (1987)

Yellow-footed marsupial “mouse”/Antechinus Flat-skulled marsupial “mouse”/Planigale Ningauis Fat-tailed marsupial “mouse”/Sminthopsis Kultarr Kowari Northern Quoll/Native “cat”/Tiger “cat” Eastern Quoll/Native “cat”/Tiger “cat” Spotted-tailed quoll Tasmanian devil

14 14 14 14 14 14 14 14 14 14

Rofe (1979) Rofe (1979) Rofe (1979) Rofe (1979) Rofe (1979) Rofe (1979) Rofe (1979) Rofe (1979) CRES San Diego (Unpublished) CRES San Diego (Unpublished)

Bilby/Rabbit-eared bandicoot Short-nosed bandicoot Long-nosed bandicoot

18F/19M 14 14

Martin and Hayman (1967) Rofe (1979) Rofe (1979)

Marsupial mole

20

Calaby et al. (1974)

Family Phalangeridae Trichosurus vulpecula Family Potoroidae Potorous tridactylus Family Macropodidae Thylogale billardierii Thylogale thetis Petrogale xanthopus celeris Petrogale xanthopus xanthopus Petrogale lateralis lateralis Petrogale penicillata Petrogale concinna

35 36

Order Diprotodontia

Family Phascolarctidae Phascolarctos cinereus Family Vombatidae Vombatus ursinus Lasiorhinus latifrons

32 33 34 35

Order Notoryctemorphia

Family Notoryctidae Notoryctes typhlops

22 23 24 25 26 27 28 29 30 30 31

Order Peramelemorphia

Family Peramelidae Macrotis (Thylacomys) lagotis Isoodon obesulus Perameles nasuta

20 21

Order Dasyuromorphia

Family Dasyuridae Antechinus flavipes Planigale maculata Ningaui sp Sminthopsis crassicaudata Antechinomys laniger Dasyuroides byrnei Dasyurus hallucatus Dasyurus viverrinus Dasyurus maculatus Sarcophilus harrisii

19

Koala

16

CRES San Diego (Unpublished)

40

Common wombat/Coarse-haired wombat Northern hairy-nosed wombat/Soft-furred wombat

14 14

Rofe (1979) Rofe (1979)

41 42

Common brush-tailed possum

20

Rofe (1979)

43

Long-nosed potoroo

12

Graphodatsky et al. (Unpublished)

44

Red-bellied pademelon Red-necked pademelon Yellow-footed rock wallaby Yellow-footed rock wallaby Black-footed rock wallaby Brush-tailed rock wallaby Little rock wallaby

22 22 22 22 22 22 16

Rofe (1979) Eldridge et al. (1992a) Eldridge et al. (1992a) Eldridge et al. (1992a) Eldridge et al. (1992a) Rofe (1979) Eldridge et al. (1992b)

45 46 46 47 47 48 49

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

CONTENTS

SPECIES

COMMON NAME

Petrogale brachyotis Wallabia bicolor Macropus parma Macropus rufogriseus Macropus eugenii Macropus eugenii Macropus parryi Macropus giganteus Macropus fuliginosus Macropus robustus Macropus rufus Family Burramyidae Cercartetus concinnus Family Pseudocheiridae Pseudocheirus peregrinus Petropseudes dahli Hemibelideus lemuroides Family Petauridae Petaurus norfolcensis Family Tarsipedidae Tarsipes rostratus Family Acrobatidae Acrobates pygmaeus

vii

2N

SOURCE*

Short-eared rock wallaby Swamp wallaby Parma wallaby Red-necked wallaby Tammar wallaby Tammar wallaby Whiptail wallaby Eastern gray kangaroo Western gray kangaroo Common wallaroo Red kangaroo

20 10F/11M 16 16 16 16 16 16 16 16 20

Eldridge et al. (1992b) Rofe (1979) CRES San Diego (Unpublished) Rofe (1979) Graphodatsky et al. (Unpublished) Rofe (1979) Rofe (1979) Rofe (1979) Rofe (1979) Rofe (1979) Rofe (1979)

49 50 51 51 52 53 54 55 56 57 57

Western pygmy possum

14

Rofe (1979)

58

Common ring-tailed possum Rock ring-tailed possum Lemuriod ring-tailed possum/Greater gliding possum

20 16 20

Murray et al. (1980) Murray et al. (1980) McQuade (1984)

59 60 60

Lesser gliding possum/Squirrel glider

21

Rofe (1979)

61

Honey possum

24

Hayman (1990)

61

Pygmy gliding possum/Feather-tailed glider

14

Rofe (1979)

62

AFROTHERIA

63

Order Afrosoricida

63

Order Macroscelidea

63

Family Chrysochloridae Chlorotalpa duthieae Chlorotalpa sclateri Amblysomus julianae Amblysomus iris Amblysomus hottentotus (durban) Amblysomus hottentotus (dullstroom) Family Tenrecidae Microgale dobsoni Hemicentetes nigriceps Echinops telfairi Family Macroscelididae Macroscelides proboscideus

African golden mole African golden mole South African golden mole South African golden mole South African golden mole South African golden mole

30 30 31 34 30 36

Bronner (1995a) Bronner (1995a) Bronner (1995a) Capanna et al. (1989) Bronner (1995a) Bronner (1995a)

65 65 66 66 67 67

Long-tailed tenrec/Shrew-like tenrec Streaked Tenrec Small Madagascar hedgehog

30 38 40

Benirschke (1969) Benirschke (1969) Benirschke (1969)

68 68 69

Short-eared elephant shrew

26

CRES San Diego (Unpublished)

70 71

Order Sirenia

Family Trichechidae Trichechus manatus latirostris Trichechus manatus latirostris

Ideogram

Florida manatee Florida manatee

48 48

Gray et al. (2002) Gray et al. (2002)

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

72 73

viii

CONTENTS

SPECIES

COMMON NAME

2N

SOURCE* 74

Order Proboscidea

Family Elephantidae Elephas maximus Loxodonta africana

Asian/Indian elephant African Savanna elephant

56 56

Houck et al. (2001) Houck et al. (2001)

78

Order Hyracoidea

Family Procaviidae Procavia capensis

Hyrax/Rock Dassie

54

Fronicke (Unpublished)

79 80

Order Tubulidentata

Family Orycteropodidae Orycteropus afer

76 77

Aardvark/Ant Bear

20

Yang et al. (2003a)

80

XENARTHRA

81

Order Xenarthra

81

Family Megalonychidae Choloepus didactylus Choloepus hoffmanni Family Myrmecophagidae Myrmecophaga tridactyla Tamandua tetradactyla (longicaudata) Cyclopes didactylus Family Dasypodidae Chaetophractus villosus Euphractus sexcinctus flavimanus Zaedyus pichiy Cabassous centralis Cabassous tatouay Tolypeutes matacus Dasypus novemcinctus Dasypus septemcinctus Dasypus hybridus

Two-toed tree sloth Two-toed tree sloth/Hoffmann’s Sloth

53 50

Nash (Unpublished) Jorge et al. (1977)

83 84

Giant anteater Lesser anteater/Tamandua Silky anteater

60 54 64

CRES San Diego (Unpublished) Jorge et al. (1977) Jorge (2000)

85 86 87

Hairy armadillo/Peludos Six-banded armadillo Pichi Naked-tailed armadillo Naked-tailed armadillo La Plata three-banded armadillo Long-nosed armadillo/Nine-banded armadillo Long-nosed armadillo/Seven-banded armadillo Long-nosed armadillo

60 58 62 62 50 38 64 64 64

Jorge et al. (1977) Jorge et al. (1977) Jorge et al. (1977) Benirschke et al. (1969) Benirschke et al. (1969) CRES San Diego (Unpublished) Jorge et al. (1977) Barroso and Seuanez (1991) Jorge et al. (1977)

88 89 90 91 91 92 92 93 93

EUARCHONTOGLIRES

95

Order Scandentia

95

Family Tupaiidae Tupaia belangeri

Northern tree shrew

62

Bigoni and Stanyon (Unpublished)

98

Order Dermoptera

Family Cynocephalidae Cynocephalus volans

Colugo/Flying lemur/Kagwang

38

Rickart (2003)

99 100

Order Primates

Family Loridae Loris tardigradus Nycticebus pygmaeus Nycticebus coucang

97

Slender loris Pygmy slow loris/Cu Lan Slow loris/Cu Lan

62 50 50

CRES San Diego (Unpublished) CRES San Diego (Unpublished) Stanyon et al. (1987a)

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

105 106 107

CONTENTS

SPECIES Galago moholi Otolemur crassicaudatus Family Cheirogaleidae Microcebus murinus Family Lemuridae Eulemur macaco flavifrons Eulemur fulvus sanfordi Family Megaladapidae Lepilemur ruficaudatus Lepilemur leucopus Lepilemur septentrionalis Family Indridae Propithecus verreauxi coquereli Propithecus tattersalli Family Daubentoniidae Daubentonia madagascariensis Daubentonia madagascariensis Ideogram Family Tarsiidae Tarsius bancanus Tarsius syrichta Family Cebidae Lagothrix lagotricha Alouatta seniculus Alouatta sara Alouatta belzebul Alouatta belzebul Ideogram Alouatta caraya Pithecia pithecia Cacajao calvus rubicundus Callicebus cupreus Callicebus pallescens Callicebus personatus nigrifrous Aotus nancymaae Aotus azarae Cebus capucinus Cebus albifrons albifrons Cebus apella Cebus apella Ideogram Cebus olivaceus Saimiri sciureus Family Callitrichidae Callimico goeldii Saguinus imperator Saguinus midas Callithrix jacchus Callithrix argentata Callithrix emiliae

COMMON NAME

2N

ix

SOURCE*

Galago/Bushbaby Brown galago

38 62

Stanyon et al. (2002) Stanyon et al. (2002)

108 109

Mouse lemur

66

CRES San Diego (Unpublished)

110

Black lemur Sanford’s lemur

44 60

CRES San Diego (Unpublished) CRES San Diego (Unpublished)

110 111

Red-tailed sportive lemur White-footed sportive lemur Northern sportive lemur

20 26 34

Rumpler et al. (1985) Rumpler et al. (1985) Rumpler et al. (1985)

112 112 113

Coquerel’s sifaka Golden-crowned sifaka

48 42

CRES San Diego (Unpublished) CRES San Diego (Unpublished)

114 114

Aye-aye Aye-aye

30 30

CRES San Diego (Unpublished) Tagle et al. (1990)

115 115

Western tarsier Philippine tarsier

80 80

Poorman et al. (1985) Rickart (2003)

116 117

Stanyon et al. (2001) Stanyon et al. (1995) Stanyon et al. (1995) Consigliere et al. (1998) Consigliere et al. (1998) Bigoni and Stanyon (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) Bigoni and Stanyon (Unpublished) Stanyon et al. (2000) Nagamachi et al. (2003) F. Garcia et al. (2002b) F. Garcia et al. (2002b) Campa and Stanyon (1992)

118 118 119 120 121 122 123 124 125 126 126 127 127 128

CRES San Diego (Unpublished) Clemente et al. (1987) Clemente et al. (1987) F. Garcia et al. (2002b) Stanyon et al. (2000)

129 130 130 131 131

Bigoni and Stanyon (Unpublished) CRES San Diego (Unpublished) Bigoni and Stanyon (Unpublished) Bigoni and Stanyon (Unpublished) Canavez et al. (1996) Nagamachi et al. (1996)

132 132 133 134 134 135

Woolly monkey Red howler Bolivian red howler Red-handed howler monkey Red-handed howler monkey Black howler monkey White-faced saki Red Uakari/Uacari Coppery titi monkey White-coated titi monkey Atlantis titi monkey Nancy Ma’s night monkey/Douroucouli Azara’s night monkey/Douroucouli White-headed Capuchin monkey/Azara’s white-faced monkey/Sapajon White-fronted Capuchin monkey Tufted Capuchin monkey Tufted Capuchin monkey Weeper Capuchin Squirrel monkey

62 44F 52 50F/49M 50F/49M 50 48 46 46 50 42 54 50 54

Goeldi’s marmoset Emperor tamarin Red-handed tamarin Common marmoset Silvery marmoset Emilia’s marmoset

48F/47M 46 46 46 44 44

54 54 54 52 44

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

x

CONTENTS

SPECIES

COMMON NAME

Callithrix humeralifer Callithrix chrysoleuca Callithrix mauesi Cebuella pygmaea Leontopithecus chrysopygus Leontopithecus rosalia Family Cercopithecidae Erythrocebus patas Cercopithecus (Chlorocebus) aethiops Cercopithecus (Chlorocebus) aethiops Cercopithecus nictitans Cercopithecus ascanius schmidti Cercopithecus petaurista Cercopithecus mona campbell Cercopithecus cephus Cercopithecus diana Cercopithecus pogonias Cercopithecus l’hoesti Miopithecus talapoin Allenopithecus nigroviridis Cercocebus galeritus Cercocebus torquatus Macaca fascicularis Macaca mulatta Papio hamadryas Macaca fuscata Macaca fuscata Mandrillus sphinx Trachypithecus obscurus Pygathrix nemaeus Trachypithecus cristatus Colobus guereza Family Hylobatidae Hylobates syndactylus Hylobates concolor Hylobates concolor Ideogram Hylobates lar Ideogram Hylobates moloch Ideogram Family Pongidae Pongo pygmaeus abelii Pongo pygmaeus Ideogram Gorilla gorilla Gorilla gorilla Ideogram Pan troglodytes Pan troglodytes Ideogram Pan paniscus Family Hominidae Homo sapiens Homo sapiens Ideogram

2N

SOURCE*

44 44 44 44 46 46

Nagamachi et al. (1996) Nagamachi et al. (1996) Nagamachi et al. (1996) Bigoni and Stanyon (Unpublished) Seuanez et al. (1988) Seuanez et al. (1988)

136 137 137 138 139 139

Patas monkey 54 African Green monkey/Grivet monkey 60 African Green monkey/Grivet monkey 60 Greater spot-nosed guenon/White-nosed monkey 70 Schmidt’s spot-nosed guenon/Red-tailed monkey 66 Lesser spot-nosed guenon 66 Mona monkey 66 Mustached guenon 66 Diana monkey 58 Guenon 72 L’Hoest monkey/L’Hoest guenon 60 Talapoin 54 Allen’s swamp monkey 48 Tana river mangabey 42 Collared Mangabey/Red-capped mangabey 42 Crab-eating macaque 42 Rhesus monkey 42 Hamadryas baboon 42 Japanese macaque 42 Japanese macaque 42 Mandrill 42 Dusty leaf monkey 44 Douc langur 44 Silver leaf monkey 44F/44M Black & white colobus monkey/Kikuya 44

Bigoni and Stanyon (Unpublished) Bigoni and Stanyon (Unpublished) Finelli et al. (1999) Ponsa and Eqozcue (1981) CRES San Diego (Unpublished) Clemente et al. (1990) Clemente et al. (1990) Clemente et al. (1990) Ponsa et al. (1986) Ponsa and Eqozcue (1981) CRES San Diego (Unpublished) Ponsa et al. (1980) CRES San Diego (Unpublished) Bigoni and Stanyon (Unpublished) F. Garcia et al. (2002) Ruiz-Herrera et al. (2002) Bigoni and Stanyon (Unpublished) Bigoni and Stanyon (Unpublished) Bigoni and Stanyon (Unpublished) Weinberg et al. (1992) Ponsa et al. (Unpublished) Ponsa et al. (1983) Bigoni (1995) Bigoni et al. (1997a) Bigoni et al. (1997b)

140 141 142 143 144 145 145 146 147 147 148 149 150 151 152 152 153 153 154 155 156 156 157 158 159

Siamang Black-crested gibbon/Concolor gibbon Black-crested gibbon/Concolor gibbon White-handed gibbon Silvery gibbon

50 52 52 44 42

Ponsa et al. (Unpublished) Bigoni and Stanyon (Unpublished) Koehler et al. (1995) Jauch et al. (1992) Stanyon and Chiarelli (1983)

160 160 161 162 163

Sumatran orangutan/Orangutan Sumatran organutan/Orangutan Lowland gorilla Lowland gorilla Common chimpanzee Common chimpanzee Pygmy chimpanzee/Bonobo

48 48 48 48 48 48 48

Bigoni and Stanyon (Unpublished) Yunis and Prakash (1982) Bigoni and Stanyon (Unpublished) Yunis and Prakash (1982) Bigoni and Stanyon (Unpublished) Yunis and Prakash (1982) Bigoni and Stanyon (Unpublished)

164 165 166 167 168 169 170

Human Human

46 46

Padilla-Nash (Unpublished) Francke (1994)

171 172

Santarem marmoset Gold and white marmoset Maués marmoset Pygmy marmoset Black lion tamarin Golden lion tamarin

Ideogram

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

CONTENTS

SPECIES

COMMON NAME

2N

SOURCE* 173

Order Rodentia

Family Sciuridae Tamias sibiricus

Chipmunk

38

Sundasciurus philippinensis Spermophilus erythrogenys

Sunda tree squirrel Ground squirrel/Susliks

38 36

Spermophilus undulatus

Ground squirrel/Susliks

32

Spermophilus alaschanicus

Ground squirrel/Susliks

38

Chestnut great flying squirrel

Petaurista albiventer Family Castoridae Castor fiber Castor canadensis Family Geomyidae Thomomys talpoides Thomomys talpoides attenuatus Thomomys talpoides pygmaeus Geomys attwateri Ideogram Family Heteromyidae Dipodomys merriami Family Dipodidae Sicista napaea Family Muridae Neotoma albigula (micropus) Peromyscus eremicus Peromyscus maniculatus Peromyscus maniculatus Ideogram Peromyscus boylii Oryzomys yunganus Peromyscus truei Peromyscus truei Ideogram Oryzomys nitidus Oryzomys megacephalus-cytotype 2 Oligoryzomys flavescens Oligoryzomys microtis Amphinectomys savamis Akodon simulator Akodon toba Abrothrix (Akodon) sanborni Abrothrix (Akodon) longipilis Chroeomys (Akodon) olivaceus Eligmodontia puerulus Eligmodontia sp Maresomy (Auliscomys) boliviensis Auliscomys sublimis Loxodontomys (Auliscomys) micropus

xi

177

38

Graphodatsky and Sablina (Unpublished) Rickart (2003) Graphodatsky and Sabina (Unpublished) Graphodatsky and Sabina (Unpublished) Graphodatsky and Sabina (Unpublished) Lie and Yang (Unpublished)

Beaver Beaver

48 40

Graphodatsky (Unpublished) Graphodatsky (Unpublished)

182 183

Western pocket gopher Western pocket gopher Western pocket gopher Pocket gopher

48 48 48 70

Thaeler Jr. (1974) Thaeler Jr. (1968) Thaeler Jr. (1968) Smolen and Bickham (1994)

184 184 185 186

Kangaroo rat

52

Mascarello et al. (1974)

187

Birch mouse

42

Graphodatsky (Unpublished)

187

Mascarello et al. (1974) Radjabli et al. (Unpublished) Greenbaum et al. (1994) Greenbaum et al. (1994) Greenbaum et al. (1994) Volobouev et al. (2000a) Greenbaum et al. (1994) Greenbaum et al. (1994) Volobouev and Aniskin (2000) Volobouev and Aniskin (2000) Aniskin and Volobouev (1999) Aniskin and Volobouev (1999) Malygin et al. (1994) Aniskin et al. (1996) Aniskin et al. (1996) Gallardo (1982) Gallardo (1982) Gallardo (1982) Kelt et al. (1991) Kelt et al. (1991) Walker and Spotorno (1992) Walker and Spotorno (1992) Walker and Spotorno (1992)

188 188 189 189 190 190 191 191 192 192 193 193 194 195 195 196 196 196 197 197 198 198 198

Wood rat/Pack rat/Trade rat 52 Cactus mouse/White-footed mouse/Deer mouse 48 White-footed mouse/Deer mouse 48 White-footed mouse/Deer mouse 48 Brush mouse/White-footed mouse/Deer mouse 48 Foothill rice rat 58 Piñon mouse/White-footed mouse/Deer mouse 48 Piñon mouse/White-footed mouse/Deer mouse 48 Elegant rice rat 80 Rice rat 54 Yellow pygmy rice rat 66 Small-eared pygmy rice rat 64 Water rat 50 Grey-bellied grass mouse 36 Chako grass mouse 42 Sanborn’s grass mouse 52 Long-haired grass mouse 52 Olive grass mouse 52 Highland desert mouse 50 Highland desert mouse 32 Bolivian big-eared mouse 22 Andean big-eared mouse 28 Southern big-eared mouse 34F/32M

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

177 178 179 180 181

xii

CONTENTS

SPECIES Euneomys chinchilloides petersoni Euneomys mordax Calomyscus mystax Calomyscus urartensis Calomyscus bailwardi Phodopus sungorus sungorus Phodopus sungorus campbelli Phodopus roborovskii Cricetus cricetus Allocricetulus eversmanni Allocricetulus curtatus Cricetulus migratorius Cricetulus barabensis Cricetulus pseudogriseus Cricetulus griseus Cricetulus obscurus Cricetulus longicaudatus Tscherskia (Cricetulus) triton Mesocricetus brandti Mesocricetus brandti Mesocricetus auratus Mesocricetus auratus Ideogram Mesocricetus raddei Mesocricetus newtoni Nannospalax (Spalax) ehrenbergi Tatera nigricauda Tachyoryctes macrocephalus Tachyoryctes splendens Gerbillus rupicola Taterillus petteri Taterillus pygargus Taterillus arenarius Taterillus tranieri Taterillus congicus Taterillus sp 1 Taterillus sp 2 Clethrionomys rufocanus Clethrionomys sikotanensis Clethrionomys rutilus Clethrionomys gapperi Clethrionomys californicus Clethrionomys glareolus Clethrionomys centralis Eothenomys regulus Alticola argentatus Alticola macrotis Phenacomys intermedius Arvicola terrestris Chionomys nivalis Chionomys roberti

COMMON NAME Patagonian chinchilla mouse Patagonian chinchilla mouse Afghan mouselike hamster Urartsk mouselike hamster Mouselike hamster Dzhungarian hamster Campbell’s hamster Small desert hamster Black-bellied hamster Eversmann’s hamster Mongolian hamster Gray dwarf hamster Striped dwarf hamster Transbaikal dwarf hamster Chinese hamster Gobi hamster Lesser long-tailed hamster Greater long-tailed hamster Brandt’s hamster Brandt’s hamster Syrian golden hamster Syrian golden hamster Ciscaucasian hamster Romanian hamster Mediterranean blind mole-rat Large naked-soled gerbil Big-headed mole-rat/African mole-rat East African mole-rat Rocky gerbil/Northern pygmy gerbil Petter’s gerbil/Small naked-soled gerbil Senegal gerbil/Small naked-soled gerbil Senegal gerbil/Small naked-soled gerbil Tranier’s gerbil/Small naked-soled gerbil Congo gerbil/Small naked-soled gerbil Small naked-soled gerbil Small naked-soled gerbil Gray red-backed vole/Red-backed vole/Bank vole Sikotan vole/Red-backed vole/Bank vole Northern red-backed vole/Bank vole Southern red-backed mouse/Bank vole Western red-backed mouse/Bank vole Red-backed vole/Bank vole Tien Shan red-backed vole Royal vole/Père David’s vole/Pratt’s vole Silver high mountain vole Large-eared vole Heather vole European water vole/Bank vole European snow vole Robert’s snow vole

2N 36 42 44 32 44 28 28 34 22 26 20 22 20 24 22 20 24 28 42 44 44 44 44 38 60 40 50 48 52 19 22 35 15 54 23 37 56 56 56 56 56 56 56 56 56 56 56 36 54 54

SOURCE* Reise and Gallardo (1990) Reise and Gallardo (1990) Radjabli et al. (Unpublished) Radjabli et al. (Unpublished) Radjabli et al. (Unpublished) Radjabli et al. (Unpublished) Radjabli et al. (Unpublished) Radjabli et al. (Unpublished) Radjabli et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Radjabli et al. (Unpublished) Radjabli et al. (Unpublished) Radjabli et al. (Unpublished) Radjabli et al. (Unpublished) Radjabli et al. (Unpublished) Radjabli et al. (Unpublished) Radjabli et al. (Unpublished) Radjabli et al. (Unpublished) Popescu and DiPaolo (1980) Radjabli et al. (Unpublished) Popescu and DiPaolo (1972) Radjabli et al. (Unpublished) Radjabli et al. (Unpublished) Ivanitskaya and Nevo (1998) Qumsiyeh (1986) Aniskin et al. (1997b) Aniskin et al. (1997b) Granjon et al. (2002) Dobigny (2002) Dobigny (2002) Volobouev and Granjon (1996) Dobigny et al. (2003) Dobigny (2002) Dobigny et al. (2002) Dobigny et al. (2002) Sokolov et al. (1990) Sokolov et al. (1990) Sokolov et al. (1990) Modi (1987) Modi (1987) Sokolov et al. (1990) Sokolov et al. (1990) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Modi (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished)

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

199 199 200 201 202 203 204 205 206 207 208 209 210 211 212 213 214 215 216 217 218 219 220 221 222 222 223 224 225 226 226 227 227 228 229 230 231 232 233 234 234 235 236 237 238 239 240 241 242 243

CONTENTS

SPECIES Chionomys gud Blanfordimys afghanus Blanfordimys bucharicus Microtus juldaschi carruthersi Microtus subterraneus Microtus daghestanicus Microtus majori Microtus nasarovi Microtus schelkovnikovi Microtus pinetorum Microtus ochrogaster Microtus richardsoni Microtus longicaudus Microtus socialis Microtus socialis schiddlovskii Microtus guentheri arm Microtus guentheri philistinus Microtus arvalis arvalis Microtus rossiaemeridionalis Microtus transcaspicus Microtus kirgisorum Microtus oeconomus Microtus montebelli Microtus fortis Microtus mongolicus Microtus hyperboreus Microtus sachalinensis Microtus maximowiczii Microtus mujanensis Microtus evoronensis Microtus gregalis Microtus agrestis Microtus pennsylvanicus Microtus breweri Microtus townsendii Microtus oregoni Microtus californicus Microtus mexicanus Lagurus lagurus Microtus sp nova from Iran Neofiber alleni Dicrostonyx torquatus Lemmus sibiricus Synaptomys cooperi Prometheomys schaposchnikowi

xiii

COMMON NAME

2N

Caucasian snow vole Afghan vole Bucharian vole Juniper vole/Vole/Meadow mouse European pine vole/Vole/Meadow mouse Daghestan pine vole/Vole/Meadow mouse Major’s pine vole/Vole/Meadow mouse Nasarov’s vole/Vole/Meadow mouse Schelkovnikov’s pine vole/Vole/Meadow mouse Campagnol/Sylvestre pine vole/Woodland vole/Vole/Meadow mouse Prairie vole/Vole/Meadow mouse Water vole/Vole/Meadow mouse Coronation Island vole/Long-tailed vole/Vole/Meadow mouse Social vole/Vole/Meadow mouse Schiddlovskii’s social vole/Vole/Meadow mouse Günther’s vole/Vole/Meadow mouse Günther’s vole/Vole/Meadow mouse Common vole/Vole/Meadow mouse Southern vole/Vole/Meadow mouse Transcaspian vole/Vole/Meadow mouse Tien Shan vole/Vole/Meadow mouse Tundra vole/Vole/Meadow mouse Japanese grass vole/Vole/Meadow mouse Reed vole Mongolian vole/Vole/Meadow mouse North Siberian vole/Vole/Meadow mouse Sakhalin vole/Vole/Meadow mouse Maximowicz’s vole/Vole/Meadow mouse Muisk vole/Vole/Meadow mouse Evoronsk vole/Vole/Meadow mouse Narrow-headed vole/Vole/Meadow mouse Field vole/Vole/Meadow mouse Campagnol des champs/Meteoro de prado/Vole/Meadow mouse Beach vole/Vole/Meadow mouse Townsend’s vole/Vole/Meadow mouse Creeping vole/Vole/Meadow mouse Amargosa vole/California vole/Vole/ Meadow mouse Mexican vole/Vole/Meadow mouse Steppe lemming Vole/Meadow mouse Round-tailed muskrat/Florida water rat Collared lemming/Varying lemming True lemming Bog lemming Long-clawed Mole-vole

54 58 48 54 54 54 54 42 54 62

Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Modi (1987)

244 245 246 247 248 249 250 251 252 253

54 56 64

Modi (1987) Modi (1987) Modi (1987)

253 254 254

62 60 60 54 46 54 52 54 30 30 52 50 50 50 44 38 40 36 50 46

Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Modi (1978) Sablina et al. (Unpublished) Kovalskaya et al. (1991) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Modi (1987) Modi (1987)

255 256 256 257 258 259 260 261 262 262 263 264 264 265 266 266 267 268 269 269

Modi (1987) Modi (1987) Modi (1987) Modi (1987)

270 270 271 271

Modi (1987) Modi (1987) Sablina et al. (Unpublished) Modi (1987) Modi (1987) Modi (1987) Modi (1987) Sablina et al. (Unpublished)

272 272 273 274 274 275 275 276

46 50 17–18 53 48 56 54 52 30 50 50 56

SOURCE*

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

xiv

CONTENTS

SPECIES

COMMON NAME

2N

Ellobius talpinus

Northern mole-vole/Mole-lemming

50

Ellobius fuscocapillus

Southern mole-vole/Mole-lemming

36

Ellobius lutescens Apodemus peninsulae

Transcaucasian mole-vole/Mole-lemming Korean wood mouse/Field mouse/Old world wood mouse Long-tailed field mouse/Field mouse/Old world wood mouse House rat/Black rat/Roof rat

Apodemus sylvaticus Rattus rattus Rattus norvegicus Rattus flavipectus Phloeomys cumingi Crunomys suncoides Archboldomys luzonensis Celaenomys silaceus Nesokia indica Leopoldamys edwardsi Mastomys natalensis Mastomys erythroleucus-cytotype 3 Mastomys awashensis Mastomys huberti Mastomys kollmanspergeri Myomys daltoni Stenocephalemys albocaudata Stenocephalemys griseicauda Stenocephalemys albipes Lophuromys melanonyx Lophuromys brevicaudus Lophuromys chrysopus Dasymys incomtus Dasymys rufulus Arvicanthis niloticus-cytotype 1A Arvicanthis niloticus-cytotype 1B Arvicanthis ansorgei Arvicanthis rufinus Arvicanthis sp. Mus musculus Mus musculus Ideogram Mus macedonicus Mus mahomet Family Pedetidae Pedetes capensis

Norway rat/Brown rat Buff-bellied rat Slender-tailed cloud rat Philippine rat/Sulawesian shrew rat Mt. Isaroq shrew rat Luzon striped rat Short-tailed bandicoot rat/Pest rat Long-tailed giant rat Multimammate rat Guinea multimammate rat Multimammate rat Multimammate rat Kollmansperger’s multimammate rat Dalton’s mouse Ethiopian narrow-headed rat Gray-tailed narrow-headed rat Ethiopian narrow-headed rat Black clawed brush-furred rat/Brush-furred mouse Short-tailed brush-furred rat/Brush-furred mouse Brush-furred rat/Brush-furred mouse African marsh rat/Shaggy swamp rat West African shaggy rat/Shaggy swamp rat African grass rat/Unstriped grass rat/Kusu rat African grass rat/Unstriped grass rat/Kusu rat African grass rat/Unstriped grass rat/Kusu rat African grass rat/Unstriped grass rat/Kusu rat African grass rat/Unstriped grass rat/Kusu rat House mouse House mouse Macedonian mouse Mahomet mouse Springhare/Springhaas

SOURCE* Perelman and Graphodatsky (Unpublished) Perelman and Graphodatsky (Unpublished) Perelman et al. (Unpublished) Sablina et al. (Unpublished)

279 279

48

Stanyon et al. (2004)

280

38

281

42 46 44 36 26 44 42 44 32 38 32 32 38 36 54 54 46 60

Sablina and Graphodatsky (Unpublished) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Rickart and Heany (2002) Rickart and Heany (2002) Rickart and Heany (2002) Rickart and Heany (2002) Sablina et al. (Unpublished) Sablina et al. (Unpublished) Volobouev et al. (2002a) Volobouev et al. (2002a) Volobouev et al. (2002a) Volobouev et al. (2002a) Volobouev et al. (2002a) Volobouev et al. (2002a) Lavrenchenko et al. (2001) Lavrenchenko et al. (2001) Lavrenchenko et al. (2001) Aniskin et al. (1997a)

282 283 284 284 285 285 286 287 288 289 290 291 292 293 294 295 296 297

68

Aniskin et al. (1997a)

298

54 36 36+B’s 62 62 62 62 58 40 40 40 36

Aniskin et al. (1997a) Volobouev et al. (2000) Volobouev et al. (2000) Volobouev et al. (2002b) Volobouev et al. (2002b) Volobouev et al. (2002b) Volobouev et al. (2002b) Volobouev et al. (2002b) Stanyon (Unpublished) Adler (1994) Ivanitskaya et al. (1996) Aniskin et al. (1998)

299 300 301 302 303 304 305 306 307 308 309 309

38

Biltueva and Graphodatsky (Unpublished)

310

17 48/+B’s

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

277 278

CONTENTS

SPECIES Family Ctenodactylidae Pectinator spekei Massoutiera mzabi Ctenodactylus gundi Ctenodactylus gundi Ctenodactylus vali

COMMON NAME

2N

xv

SOURCE*

Speke’s pectinator Massoutiera Atlas gundi Atlas gundi Sahara gundi

40 36 40 40 40

George (1979) George (1979) George (1979) George (1979) George (1979)

311 311 312 312 313

Family Myoxidae Muscardinus avellanarius Myoxus glis Dryomys nitedula Eliomys quercinus Myomimus personatus

Common dormouse/Hazel mouse Fat dormouse/Edible dormouse Forest dormouse Garden dormouse Mouselike dormouse

46 62 48 48 44

Graphodatsky (Unpublished) Graphodatsky (Unpublished) Graphodatsky (Unpublished) Graphodatsky (Unpublished) Graphodatsky (Unpublished)

314 315 316 317 318

Family Bathyergidae Georychus capensis Bathyergus janetta Cryptomys mechowi Cryptomys hottentotus damarensis

Cape mole-rat Dune mole-rat Giant mole-rat/Common mole-rat Common mole-rat

54 54 40 78

Nevo et al. (1986) Nevo et al. (1986) Macholan et al. (1993) Nevo et al. (1986)

319 319 320 321

Family Chinchillidae Lagostomus maximus

Plains viscacha

56

Graphodatskaya and Graphodatsky (Unpublished)

322

Family Dinomyidae Dinomys branickii

Pacarana

58

CRES San Diego (Unpublished)

323

Family Caviidae Cavia porcellus (cobaya)

Guinea pig/Cavies

64

Graphodatsky and Sablina (Unpublished)

324

Family Hydrochaeridae Hydrochaeris hydrochaeris

Capybara

66

CRES San Diego (Unpublished)

325

Family Ctenomyidae Ctenomys rionegrensis Ctenomys talarum Ctenomys maulinus brunneus Ctenomys robustus Ctenomys maulinus maulinus Ctenomys magellanicus magellanicus Ctenomys opimus Ctenomys fulvus

Tuco-tuco Tuco-tuco Tuco-tuco Tuco-tuco Tuco-tuco Tuco-tuco Tuco-tuco Tuco-tuco

50 48 26 26 26 34 26 26

L. Garcia et al. (2002) L. Garcia et al. (2002) Gallardo (1979) Gallardo (1979) Gallardo (1979) Gallardo (1979) Gallardo (1979) Gallardo (1979)

325 326 326 326 327 327 327 328

Family Octodontidae Octodon lunatus Octodon degus Tympanoctomys barrerae Tympanoctomys barrerae Spalacopus cyanus Aconaemys fuscus Aconaemys sagei Aconaemys porteri

Degus Degus Viscacha rat Viscacha rat Coruro Rock rat Rock rat Rock rat

78 58 102 102 58 56 54 58

Spotorno et al. (1995) Spotorno et al. (1995) Gallardo et al. (2004) Gallardo et al. (2004) Spotorno et al. (1995) Gallardo and Mondaca (2002) Gallardo and Mondaca (2002) Gallardo and Mondaca (2002)

328 329 330 331 332 333 333 333

Ideogram

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

xvi

CONTENTS

SPECIES Family Abrocomidae Abrocoma bennetti Family Echimyidae Proechimys brevicauda-cytotype 1 Proechimys steerei Proechimys simonsi Proechimys sp.1 Proechimys sp.2 Lonchotrix emiliae Isothrix sinnamariensis Dactylomys dactylinus Family Myocastoridae Myocastor coypus

COMMON NAME

2N

SOURCE*

Chinchilla rat/Chinchillone

64

Spotorno et al. (1995)

334

Huallaga spiny rat/Casiragua Steere’s spiny rat/Casiragua Simon’s spiny rat/Casiragua Terrestrial spiny rat/Casiragua Terrestrial spiny rat/Casiragua Tuft-tailed spiny tree rat French Guiana brush-tailed rat/Toros Amazon bamboo rat/Coro-coro

28 24 32 30 34 62 28 96

Aniskin (1994b) Aniskin (1994b) Aniskin (1994b) Aniskin (1994b) Aniskin (1994b) Aniskin (1994b) Vié et al. (1996) Aniskin (1993)

335 336 337 338 338 339 339 340

Nutria/Coypu

42

Biltueva and Graphodatsky (Unpublished)

341 342

Order Lagomorpha

Family Ochotonidae Ochotona princeps Ochotona hyperborea

American pika/Mouse hare/Conie Northern pika/Mouse hare/Conie

68 40

Stock (1976) Perelman and Graphodatsky (Unpublished)

345 346

Family Leporidae Pronolagus rupestris Bunolagus monticularis Romerolagus diazi Brachylagus idahoensis Sylvilagus aquaticus Sylvilagus palustris Sylvilagus transitionalis Sylvilagus floridanus Sylvilagus nuttallii Sylvilagus audubonii Lepus californicus Lepus saxatilis Oryctolagus cuniculus Oryctolagus cuniculus Ideogram

Red rabbit Bushman rabbit Volcano rabbit Pygmy rabbit Swamp rabbit/Cottontail Marsh rabbit/Cottontail New England cottontail Eastern cottontail Mountain cottontail Desert cottontail Black-tailed jack rabbit/Hare Jack rabbit/Hare Domestic rabbit/Old world rabbit Domestic rabbit/Old world rabbit

42 44 48 44 38 38 46 42 42 42 48 48 44 44

Robinson et al. (2002) Robinson et al. (2002) Robinson et al. (2002) Robinson et al. (2002) Robinson et al. (1983) Robinson et al. (1983) Robinson et al. (1983) Robinson et al. (1983) Robinson et al. (1984) Robinson et al. (1984) Robinson et al. (2002) Robinson (1980) Robinson et al. (2002) Hayes et al. (2002)

346 347 348 349 350 350 351 351 352 352 353 354 354 355

LAURASIATHERIA

357

Order Eulipotyphla

357

Family Erinaceidae Podogymnura truei Erinaceus europaeus

Philippine Gymnure/Philippine wood shrew Western Eurasian hedgehog

40 48

Erinaceus concolor Erinaceus amurensis

Eastern Eurasian hedgehog Eurasian hedgehog/Amur hedgehog

48 48

Paraechinus hypomelas

Brandt’s hedgehog/Desert hedgehog

48

Rickart (2003) Rabjabli and Graphodatsky (Unpublished) Sokolov et al. (1991) Radjabli and Graphodatsky (Unpublished) Radjabli and Graphodatsky (Unpublished)

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

360 360 361 362 363

CONTENTS

SPECIES

COMMON NAME

2N

xvii

SOURCE*

Hemiechinus auritus

Long-eared desert hedgehog

48

Mesechinus (Hemiechinus) dauricus

Daurian hedgehog

48

Hispaniolan Solenodon

34

Benirschke (1969)

366

Eurasian pygmy shrew/Long-tailed shrew Laxmann’s shrew/Long-tailed shrew Radde’s shrew/Long-tailed shrew Eurasian shrew/Long-tailed shrew Eurasian shrew/Long-tailed shrew Lagranja shrew/Long-tailed shrew Tundra shrew Southern short-tailed shrew Southern short-tailed shrew Southern short-tailed shrew Elliot’s short-tailed shrew Old world water shrew

42 42 36 21 21 36 30 46 37 50 19 52

366 367 367 368 368 369 369 370 370 371 371 372

Crocidura suaveolens

Lesser shrew/White-toothed shrew

40

Crocidura horsfieldii watasei

Horsfield’s shrew/White-toothed shrew

26

Pale gray shrew/White-toothed shrew Siberian shrew/White-toothed shrew Musk shrew/Pygmy shrew Dsinezumi shrew/White-toothed shrew

22 40 40 40

Perelman et al. (Unpublished) Perelman et al. (Unpublished) Perelman et al. (Unpublished) Perelman et al. (Unpublished) Serov et al. (1998) Perelman et al. (Unpublished) Aniskin and Volobouev (1980) George et al. (1982) George et al. (1982) George et al. (1982) George et al. (1982) Perelman and Graphodatsky (Unpublished) Perelman and Graphodatsky (Unpublished) Perelman and Graphodatsky (Unpublished) Graphodatsky (Unpublished) Graphodatsky (Unpublished) Perelman et al. (Unpublished) Perelman et al. (Unpublished)

Old world mole Siberian mole/Old world mole Hairy-tailed mole

34 34 34

Benirschke (1969) Graphodatsky (Unpublished) Benirschke (1969)

376 377 377

Family Solenodontidae Solenodon paradoxus Family Soricidae Sorex minutus Sorex caecutiens Sorex raddei Sorex araneus Sorex araneus Ideogram Sorex granarius Sorex tundrensis Blarina carolinensis Blarina carolinensis Blarina carolinensis peninsulae Blarina hylophaga Neomys fodiens

Crocidura pergrisea Crocidura sibirica Suncus murinus Crocidura dsinezumi Family Talpidae Talpa europaea europaea Talpa altaica Parascalops breweri

Radjabli and Graphodatsky (Unpublished) Radjabli and Graphodatsky (Unpublished)

365

373 373 374 374 375 376

378

Order Chiroptera

Family Pteropodidae Rousettus aegyptiacus Rousettus leschenaulti Rousettus lanosus Rousettus (Lissonycteris) angolensis Myonycteris torquata Pteropus rodricensis Pteropus giganteus Pteropus lylei Hypsignathus monstrosus Epomops franqueti Micropteropus pusillus Scotonycteris ophiodon Otopteropus cartilagonodus Alionycteris paucidentata Eonycteris robusta Megaloglossus woermanni

364

Rousette fruit bat Rousette fruit bat Rousette fruit bat Rousette fruit bat Little-collared fruit bat Rodriguez flying fox Indian flying fox Flying fox Hammer-headed fruit bat Epauleted bat Dwarf epauleted bat Long-haired tailless flying fox Long-haired flying fox Dawn bat African long-tongued fruit bat

36 36 36 36 36 38 38 40 36 36 35 34 48 36 36 36

Haiduk et al. (1981) Sreepada and Gururaj (1995) Teeling and Stanyon (Unpublished) Haiduk et al. (1981) Haiduk et al. (1981) CRES San Diego (Unpublished) Choudhury and Jena (1995) Hood et al. (1988) Haiduk et al. (1981) Haiduk et al. (1981) Haiduk et al. (1981) Haiduk et al. (1981) Rickart et al. (1999) Rickart et al. (1999) Rickart et al. (1999) Haiduk et al. (1981)

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

381 381 382 382 383 383 384 384 385 385 386 386 387 387 388 388

xviii

CONTENTS

SPECIES Family Rhinopomatidae Rhinopoma microphyllum Rhinopoma hardwickei Family Emballonuridae Taphozous longimanus Taphozous melanopogon Taphozous nudiventris Taphozous saccolaimus Family Megadermatidae Megaderma spasma Megaderma lyra Family Rhinolophidae Rhinolophus rouxi Rhinolophus ferrumequinum Family Hipposideridae Hipposideros pomona Hipposideros ater Hipposideros fulvus Hipposideros fulvus pallidus Hipposideros cineraceus Hipposideros lankadiva Hipposideros speoris Family Mormoopidae Pteronotus parnellii Ideogram Pteronotus personatus Pteronotus macleayii Pteronotus gymnonotus Mormoops blainvillii Family Noctilionidae Noctilio albiventris Family Phyllostomidae Desmodus rotundus Diaemus youngi Diphylla ecaudata Macrotus waterhousii Micronycteris megalotis Micronycteris schmidtorum Micronycteris minuta Micronycteris hirsuta Micronycteris brachyotis Micronycteris nicefori Vampyrum spectrum Trachops cirrhosus

COMMON NAME

2N

SOURCE*

Mouse-tailed bat/Long-tailed bat Mouse-tailed bat/Long-tailed bat

42 36

Qumsiyeh (1988) Qumsiyeh (1988)

389 389

Tomb bat Tomb bat Tomb bat Tomb bat

42 42 42 44

Sreepada et al. (1995) Sreepada et al. (1995) Sreepada et al. (1995) Sreepada et al. (1995)

390 390 390 391

Asian false vampire bat Asian false vampire bat

38 54

Morielle-Versute et al. (1992) Choudhury and Mohanty (1993)

391 392

Horseshoe bat Horseshoe bat

56 56

Zima et al. (1992) 392 Teeling and Stanyon (Unpublished) 393

Old world leaf-nosed bat Old world leaf-nosed bat Old world leaf-nosed bat Old world leaf-nosed bat Old world leaf-nosed bat Old world leaf-nosed bat Old world leaf-nosed bat

32 32 32 32 32 32 32

Sreepada et al. (1993) Sreepada et al. (1993) Sreepada et al. (1993) Choudhury and Patro (1993) Sreepada et al. (1993) Sreepada et al. (1993) Sreepada et al. (1993)

394 394 394 395 395 395 396

Moustached bat/naked-backed bat/leaflipped bat Moustached bat/naked-backed bat/leaflipped bat Moustached bat/naked-backed bat/leaflipped bat Moustached bat/naked-backed bat/leaflipped bat Leaf-chinned bat

38

Sites et al. (1981)

397

38

Sites et al. (1981)

397

38

Sites et al. (1981)

398

38

Sites et al. (1981)

398

38

Sites et al. (1981)

399

Bulldog bat/fisherman’s bat

34

Varella-Garcia et al. (1989)

399

Common vampire bat White-winged vampire bat Hairy-legged vampire bat Big-eared bat Little big-eared bat Little big-eared bat Little big-eared bat Little big-eared bat Little big-eared bat Little big-eared bat Linnaeus’s false vampire bat/Spectral vampire bat Frog-eating bat

28 32 32 46 40 38 28 30 32 28 30

Varella-Garcia et al. (1989) Varella-Garcia et al. (1989) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979)

400 401 401 402 402 403 403 404 404 404 405

30

Baker et al. (1979)

405

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

CONTENTS

SPECIES Chrotopterus auritus Phylloderma stenops Phyllostomus discolor Phyllostomus hastatus Tonatia bidens Mimon crenulatum Glossophaga soricina Monophyllus redmani Lonchophylla robusta Lonchophylla thomasi Lionycteris spurrelli Anoura caudifera Anoura cultrata Lichonycteris obscura Hylonycteris underwoodi Choeronycteris mexicana Choeroniscus intermedius Erophylla sezekorni Choeroniscus godmani Phyllonycteris poeyi Carollia castanea Carollia brevicauda Carollia perspicillata Rhinophylla pumilio Rhinophylla fischerae Sturnira lilium Sturnira mordax Sturnira erythromos Uroderma magnirostrum Platyrrhinus vittatus Platyrrhinus vittatus lineatus Vampyressa pusilla Vampyressa nymphaea Vampyressa brocki Chiroderma villosum Chiroderma doriae Mesophylla macconnelli Chiroderma improvisum Ectophylla alba Enchisthenes hartii Artibeus jamaicensis Ardops nichollsi Artibeus lituratus Phyllops haitiensis Ariteus flavescens Ametrida centurio Sphaeronycteris toxophyllum Family Mystacinidae Mystacina tuberculata

COMMON NAME

2N

xix

SOURCE*

Peter’s woolly false vampire bat 28 Peter’s spear-nosed bat 32 Spear-nosed bat 32 Spear-nosed bat 30 Round-eared bat 16 Gray’s spear-nosed bat 32 Long-tongued bat 32 Leach’s single leaf bat 32 Saussure’s long-nosed bat 28 Saussure’s long-nosed bat 30 Chestnut long-nosed bat 28 Geoffroy’s long-nosed bat 30 Geoffroy’s long-nosed bat 30 Dark long-tongued bat 24 Underwood’s long-tongued bat 16 Mexican long-nosed bat/Hog-nosed bat 16 Godman’s long-nosed bat 20 Brown flower bat 32 Godman’s long-nosed bat 19 Cuban flower bat 32 Short-tailed leaf-nosed bat 22 Short-tailed leaf-nosed bat 20F/21M Short-tailed leaf-nosed bat 20F/21M Dwarf little fruit bat 36 Fischer’s little fruit bat 34 Yellow shouldered bat/American epauleted bat 30 Yellow shouldered bat/American epauleted bat 30 Yellow shouldered bat/American epauleted bat 30 Tent-building bat 38 White-lined bat 30 White-lined bat 30 Yellow-eared bat 18 Yellow-eared bat 26 Yellow-eared bat 24 Big-eyed bat/White-lined bat 26 Big-eyed bat/White-lined bat 26 Little yellow-faced bat 21 Big-eyed bat/White-lined bat 26 White bat 30 Hart’s little fruit bat 31 Neotropical fruit bat 30 Tree bat 31 Neotropical fruit bat 31 Falcate-winged bat 31 Jamaican fig-eating bat 31 Little white-shouldered bat 31 Red fruit bat/Visored bat 28

Morielle-Versute et al. (1992) Baker et al. (1979) Baker et al. (1979) Varella-Garcia et al. (1989) Baker et al. (1979) Baker et al. (1979) Varella-Garcia et al. (1989) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979) Varella-Garcia et al. (1989) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979) Stock (1975) Baker et al. (1979) Hsu et al. (1968) Baker et al. (1979) Stock (1975) Stock (1975) Teeling and Stanyon (Unpublished) Baker et al. (1979) Baker et al. (1979) Varella-Garcia et al. (1989) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979) Varella-Garcia et al. (1989) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979) Varella-Garcia et al. (1989) Varella-Garcia et al. (1989) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979) Varella-Garcia et al. (1989) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979) Baker et al. (1979)

406 406 407 407 408 408 409 409 410 410 410 411 411 412 412 412 413 413 413 414 414 414 415 415 416 416 417 417 417 418 418 419 419 419 420 421 422 422 422 423 423 423 424 425 425 425 426

New Zealand short-tailed bat

Bickham et al. (1980)

426

36

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

xx

CONTENTS

SPECIES Family Natalidae Natalus major Family Vespertilionidae Myotis keenii Myotis thysanodes Myotis mystacina Myotis sodalis Myotis nigricans Myotis nigricans Ideogram Pipistrellus pipistrellus Pipistrellus ceylonicus Pipistrellus javanicus Pipistrellus coromandra Pipistrellus (Hypsugo) crassulus Pipistrellus kuhli Pipistrellus (Vespadelus) vulturnus Pipistrellus (Vespadelus) sagittula Pipistrellus stenopterus Pipistrellus (Falsistrellus) tasmaniensis Pipistrellus mimus Pipistrellus (Hypsugo) eisentrauti Scotozous dormeri Hesperoptenus blandfordi Chalinolobus morio Tylonycteris pachypus Tylonycteris robustula Scotorepens balstoni Eptesicus bottae Eptesicus fuscus Scotophilus kuhlii Scotophilus heathi Corynorhinus townsendii Lasiurus borealis Idionycteris phyllotis Euderma maculatum Murina cyclotis Nyctophilus gouldi Family Molossidae Molossops temminckii Molossops abrasus Nyctinomops laticaudatus Eumops perotis Eumops glaucinus Molossus molossus Molossus rufus(ater)

COMMON NAME

2N

SOURCE*

Funnel-eared bat

36

Hoyt and Baker (1980)

427

Little brown bat Little brown bat Little brown bat Little brown bat/Indiana bat Little brown bat Little brown bat Pipistrelle Pipistrelle Pipistrelle Pipistrelle Pipistrelle Pipistrelle Pipistrelle Pipistrelle Pipistrelle Pipistrelle Pipistrelle Pipistrelle Dormer’s bat Blandford’s bat Chocolate bat/Lobe-lipped bat/Groove-lipped bat/Wattled bat Club-footed bat/Bamboo bat Club-footed bat/Bamboo bat Lesser broad-nosed bat Big brown bat/House bat/Serotine bat Big brown bat/House bat/Serotine bat

44 44 44 44 44 44 44 36 34 30 30 42 44 44 32 44 38 42 30 32 44

Bickham (1979) Bickham (1979) Volleth (1987) Bickham (1979) Bickham (1979) Bickham (1979) Volleth (1987) Sreepada et al. (1996) Volleth et al. (2001) Sreepada et al. (1996) Volleth et al. (2001) Volleth et al. (2001) Volleth and Tidemann (1989) Volleth and Tidemann (1989) Volleth et al. (2001) Volleth and Tidemann (1991) Volleth et al. (2001) Volleth et al. (2001) Volleth et al. (2001) Volleth et al. (2001) Volleth and Tidemann (1989)

427 427 428 428 428 429 429 430 430 430 431 431 432 432 432 433 433 434 434 435 435

32 46 30 50 50

435 436 436 437 438

House bat/Yellow bat House bat/Yellow bat Lump-nosed bat/American long-eared bat Hairy-tailed bat/Red bat Allen’s big-eared bat/Lappet-eared bat Spotted bat/Pinto bat Tube-nosed insectivora bat Australian big-eared bat

36 36 32 28 30 30 44 44

Volleth et al. (2001) Volleth et al. (2001) Volleth and Tidemann (1991) Volleth et al. (2001) Teeling and Stanyon (Unpublished) Stock (1975) Stock (1975) Stock (1983) Varella-Garcia et al. (1989) Stock (1983) Stock (1983) Rickart et al. (1999) Volleth and Tidemann (1989)

Broad-faced bat Broad-faced bat New world free-tailed bat Mastiff bat/Bonneted bat Mastiff bat/Bonneted bat Velvety free-tailed bat Velvety free-tailed bat

48 34 48 48 40 48 48

Morielle-Versute et al. (1996) Morielle-Versute et al. (1996) Morielle-Versute et al. (1996) Morielle-Versute et al. (1996) Morielle-Versute et al. (1996) Morielle-Versute et al. (1996) Morielle-Versute et al. (1996)

442 443 443 443 444 444 444

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

438 439 439 440 440 441 441 442

CONTENTS

SPECIES

COMMON NAME

2N

SOURCE* 445

Order Carnivora

Family Canidae Vulpes vulpes Vulpes vulpes Ideogram Vulpes corsac Vulpes macrotis Fennecus zerda Alopex lagopus Alopex lagopus Ideogram Urocyon cinereoargenteus Cerdocyon thous Cerdocyon thous Ideogram Nyctereutes procyonoides viverrinus Nyctereutes procyonoides viverrinus Ideogram Nyctereutes procyonoides procyonoides Atelocynus microtis Speothos venaticus Canis latrans Canis lupus Canis familiaris Canis familiaris Ideogram Chrysocyon brachyurus Otocyon megalotis Family Ursidae Tremarctos ornatus Tremarctos ornatus Ideogram Ursus thibetanus Ursus americanus Ursus arctos Ursus maritimus Ursus malayanus Ursus ursinus Ailuropoda melanoleuca Ailuropoda melanoleuca Ideogram Family Procyonidae Ailurus fulgens Bassariscus astutus Potos flavus Procyon lotor Procyon lotor Ideogram Bassaricyon sp. Family Mustelidae Mustela erminea Mustela nivalis

xxi

Red fox

34 + Bs

Red fox Corsac fox Kit fox Fennec fox Arctic fox Arctic fox Grey fox Crab-eating fox Crab-eating fox Japanese raccoon dog Japanese raccoon dog Chinese raccoon dog Small-eared dog/Round-eared dog Bush dog Coyote Grey wolf Domestic dog Domestic dog Maned wolf Bat-eared fox

34 + Bs 36 50 64 50 50 66 74 74 38 + Bs 38 + Bs 54 + Bs 74 74 78 78 78 78 76 72

Graphodatsky and Beklemisheva (Unpublished) Rubtsova (1998) CRES San Diego (Unpublished) Wayne et al. (1987a) Graphodatsky (Unpublished) Nash et al. (2001) Nash et al. (2001) Wayne et al. (1987a) Nash et al. (2001) Nash et al. (2001) Nash et al. (2001) Nash et al. (2001) Graphodatsky et al. (Unpublished) Wurster-Hill and Centerwall (1982) Wayne et al. (1987a) Wurster-Hill and Centerwall (1982) Wurster-Hill and Centerwall (1982) Graphodatsky et al. (2000a) Graphodatsky et al. (2000b) Wayne et al. (1987a) Wayne et al. (1987a)

451 452 453 453 454 455 456 457 458 459 460 461 462 463 464 465 466 467 468 469 470

Spectacled bear Spectacled bear Asiatic black bear American black bear Brown bear Polar bear Sun bear Sloth bear Giant panda Giant panda

52 52 74 74 74 74 74 74 42 42

Nash (Unpublished) Nash (Unpublished) Nash and O’Brien (1987) Nash and O’Brien (1987) Nash and O’Brien (1987) Nash and O’Brien (1987) Nash and O’Brien (1987) Nash and O’Brien (1987) Nash (Unpublished) Nash (Unpublished)

471 472 473 474 475 476 477 478 479 480

Lesser panda/Red panda Ringtail/Cacomistle Kinkajou Raccoon Raccoon Olingo

36 38 38 38 38 38

Nash (Unpublished) Nash (Unpublished) CRES San Diego (Unpublished) Stanyon (Unpublished) Stanyon et al. (1993) Wurster-Hill and Gray (1975)

481 481 482 482 483 484

Ermine/Stoat Least weasel

44 42

484 485

Mustela altaica

Mountain weasel

44

Mustela lutreola

European mink

38

Graphodatsky (Unpublished) Perelman and Graphodatsky (Unpublished) Perelman and Graphodatsky (Unpublished) Graphodatsky (Unpublished)

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

486 487

xxii

CONTENTS

SPECIES

COMMON NAME

2N

SOURCE*

38 38 30 30 40 40 38 38 38 38 38 38 40 38 38 38 42 40 44 38 60 64 36

Graphodatsky (Unpublished) Graphodatsky (Unpublished) Nash (Unpublished) Serov (1998) Graphodatsky (Unpublished) Graphodatsky (Unpublished) Graphodatsky (Unpublished) Nie et al. (2002) Graphodatsky (Unpublished) Graphodatsky (Unpublished) Graphodatsky (Unpublished) Perelman et al. (Unpublished) Nie et al. (2002) Wurster-Hill and Centerwall (1982) Wurster-Hill and Centerwall (1982) Perelman et al. (Unpublished) Graphodatsky (Unpublished) Wurster-Hill and Centerwall (1982) Nie et al. (2002) Wurster-Hill and Centerwall (1982) Benirschke (1969) Benirschke (1969) Graphodatsky (Unpublished)

487 488 488 489 489 490 491 491 492 493 493 494 494 495 495 496 497 497 498 498 499 499 500

Wurster-Hill and Gray (1975)

500 501 501 502 502 503 503 504

Mustela sibirica Mustela sibirica itatsi Mustela vison Mustela vison Ideogram Mustela putorius putorius Mustela putorius furo Mustela eversmanni Martes foina Vormela peregusna Martes martes Martes zibellina Martes melampus Martes flavigula Eira barbara Galictis vittata Ictonyx striatus Gulo gulo Mellivora capensis Meles meles Melogale sp. Spilogale gracilis latifrons Spilogale putorius interrupta Lutra lutra Family Nandiniidae Nandinia binotata Family Viverridae Genetta genetta Genetta tigrina Prionodon linsang Viverricula indica Paradoxurus hermaphroditus Arctictis binturong Paguma larvata

Siberian weasel Japanese weasel American mink American mink Wild polecat Domesticated ferret Steppe polecat Beech marten Marbled polecat Pine marten Sable Japanese marten Yellow-throated marten Tayra Allamand’s grisón/Greater grisón Striped polecat/Zorilla Wolverine Ratel/Honey badger Old world badger Ferret badger Spotted skunk Spotted skunk European river otter

Small spotted genet Large spotted genet Banded linsang/Oriental linsang Lesser oriental civet/Rasse Palm civet Binturong Masked palm civet

54 54 34 38 42 42 44

Hemigalus derbyanus Fossa fossana Family Herpestidae Galidia elegans Herpestes javanicus Helogale parvula Bdeogale sp. Atilax paludinosus Cryptoprocta ferox Family Hyaenidae Parahyaena brunnea

Banded palm civet Malagasy civet

42 42

CRES San Diego (Unpublished) CRES San Diego (Unpublished) Wurster-Hill and Gray (1975) Wurster-Hill and Gray (1975) Wurster-Hill and Gray (1975) Wurster-Hill and Gray (1975) Graphodatsky and Beklemisheva (Unpublished) Wurster-Hill and Gray (1975) Nash (Unpublished)

Malagasy ring-tailed mongoose Java mongoose Dwarf mongoose Black-legged mongoose African marsh mongoose/Water mongoose Fossa

44 36 36 36 35 42

Wurster-Hill and Gray (1975) Yang and Nie (Unpublished) Nash (Unpublished) Wurster-Hill and Gray (1975) Wurster-Hill and Gray (1975) CRES San Diego (Unpublished)

506 506 507 507 508 508

Brown hyena

40

509

Spotted hyena

40

Perelman and Graphodatsky (Unpublished) Nash (Unpublished)

Crocuta crocuta

African Palm Civet

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

505 505

510

CONTENTS

SPECIES Family Felidae Leopardus pardalis Leopardus tigrinus Lynchailurus colocolo Oncifelis geoffroyi Felis catus Felis catus Ideogram Felis chaus Felis margarita Felis nigripes Panthera leo Panthera tigris altaica Neofelis nebulosa diardi Neofelis nebulosa Puma concolor Acinonyx jubatus Herpailurus yagouaroundi Lynx lynx Prionailurus bengalensis Prionailurus viverrinus Caracal caracal Profelis temmincki Leptailurus serval Octocolobus manul Family Otariidae Callorhinus ursinus Arctocephalus pusillus Zalophus californianus Eumetopias jubatus Family Odobenidae Odobenus rosmarus Family Phocidae Monachus schauinslandi Leptonychotes weddelli Mirounga angustirostris Erignathus barbatus Cystophora cristata Halichoerus grypus Phoca groenlandica Phoca hispida Phoca fasciata Phoca vitulina Phoca vitulina Ideogram

COMMON NAME

2N

SOURCE*

Ocelot Tiger cat/Little spotted cat Pampas cat Geoffroy’s cat Domestic cat Domestic cat Jungle cat Sand cat Black-footed cat Lion Siberian tiger Bornean clouded leopard Mainland clouded leopard Puma/Cougar/Mountain lion Cheetah Jaguarundi Eurasian lynx Asian leopard cat Fishing cat Caracal Asian golden cat Serval Pallas’s cat

36 38 38 36 38 38 38 38 38 38 38 38 38 38 38 38 38 38 38 38 38 38 38

Wurster-Hill and Gray (1975) Wurster-Hill and Centerwall (1982) Wurster-Hill and Gray (1973) Nash (Unpublished) Nash (Unpublished) Cho et al. (1997) Wurster-Hill and Gray (1973) Wurster-Hill and Gray (1975) Nash (Unpublished) Nash (Unpublished) CRES San Diego (Unpublished) Nash (Unpublished) Nash (Unpublished) Wurster-Hill and Gray (1973) Nash (Unpublished) Wurster-Hill and Gray (1973) Yang and Li (Unpublished) Wurster-Hill and Gray (1973) Wurster-Hill and Gray (1973) Wurster-Hill and Gray (1973) Wurster-Hill and Gray (1973) Wurster-Hill and Centerwall (1982) Wurster-Hill and Gray (1975)

511 511 512 513 514 515 516 516 517 518 519 520 521 522 523 524 524 525 526 527 527 528 529

Northern fur seal South African fur seal/Southern fur seal California sea lion Northern sea lion/Steller sea lion

36 36 36 36

Arnason (1977) Arnason (1977) Arnason (1974b) Arnason (1974b)

530 530 531 531

Walrus

32

Arnason (1977)

532

Hawaiian monk seal Weddell seal Northern elephant seal Bearded seal Hooded seal Gray seal Harp seal Ringed seal Ribbon seal Harbor seal/Common seal Harbor seal/Common seal

34 34 34 34 34 32 32 32 32 32 32

Arnason (1974b) Arnason (1974b) Arnason (1974b) Arnason (1974b) Arnason (1974b) Arnason (1970a) Arnason (1974c) Arnason (1974c) Arnason (1977) Nash (Unpublished) Fröenicke (Unpublished)

532 533 533 534 534 535 535 536 536 537 538 539

Order Pholidota

Family Manidae Manis javanica

xxiii

Javan pangolin/Malayan pangolin/Scaly anteater

38

Nie and Yang (Unpublished)

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

540

xxiv

CONTENTS

SPECIES

COMMON NAME

2N

SOURCE* 541 541

Order Cetartiodactyla

Cetacea Family Lipotidae Lipotes vexillifer Family Monodontidae Delphinapterus leucas Family Phocoenidae Phocoena phocoena Family Delphinidae Stenella clymene Stenella attenuata Stenella dubia Delphinus delphis Lagenorhynchus albirostris Tursiops truncatus Tursiops truncatus Ideogram Orcinus orca Globicephala macrorhynchus Family Ziphiidae Mesoplodon carlhubbsi Mesoplodon europaeus Family Physeteridae Kogia breviceps Physeter macrocephalus Family Eschrichtiidae Eschrichtius robustus Family Balaenidae Balaena mysticetus Family Balaenopteridae Balaenoptera acutorostrata Balaenoptera physalus Balaenoptera borealis Balaenoptera musculus Megaptera novaeangliae

Baiji/Chinese river dolphin

44

Minrong et al. (1986)

547

Beluga whale/White whale

44

Jarrell and Arnason (1981)

548

Harbor porpoise/Common porpoise

44

Arnason (1980)

549

Atlantic spinner dolphin Pantropical spotted dolphin Spotted dolphin Common dolphin/Saddlebacked dolphin White-beaked dolphin Bottle-nosed dolphin Bottle-nosed dolphin Killer whale Pilot whale/Blackfish whale

44 44 44 44 44 44 44 44 44

Arnason (1980) Stock (1981) Arnason (1974c) Arnason (1974a) Arnason (1980) CRES San Diego (Unpublished) Bielec et al. (1997) Arnason et al. (1980) Arnason (1974a)

549 550 551 551 552 552 553 554 555

Hubbs’ beaked whale Gervais’ beaked whale

42 42

Arnason et al. (1977) Arnason et al. (1977)

555 556

Pygmy sperm whale Sperm whale

42 42

Arnason and Benirscke (1973) Arnason and Benirscke (1973)

557 558

Gray whale

44

Arnason (1981a)

559

Bowhead whale/Greenland right whale

42

Jarelle (1979)

559

Minke whale Fin whale Sei whale Blue whale Humpback whale

44 42 44 44 44

Arnason (1981a) Arnason (1974a) Nash (Unpublished) Arnason and Widegren (1989) Arnason (1974a)

560 560 561 562 563 564 564

Order Cetartiodactyla

Artiodactyla Family Suidae Sus scrofa domestica Sus scrofa domestica Ideogram Sus scrofa scrofa Sus barbatus Sus cebifrons Potamochoerus porcus Potamochoerus larvatus Phacochoerus africanus sundervallii Babyrousa babyrussa celebensis

Domestic pig/Hog/Boar Domestic pig/Hog/Boar European wild boar/Hog/Boar Bearded pig/Hog/Boar Visayan warty pig/Hog/Boar Red river hog Southern bush pig/African bush pig Southern warthog Sulawesi babirusa

38 38 36 38 34 34 34 34 38

Graphodatsky (Unpublished) Biltueva et al. (2004) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished)

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

566 567 567 568 568 569 570 571 571

CONTENTS

SPECIES Family Tayassuidae Catagonus wagneri Tayassu pecari Tayassu tajacu Family Hippopotamidae Hippopotamus amphibius Family Camelidae Camelus bactrianus Camelus dromedarius Family Tragulidae Tragulus javanicus Family Giraffidae Okapia johnstoni Giraffa camelopardalis tippelskirchi Family Moschidae Moschus moschiferus

COMMON NAME

xxv

2N

SOURCE*

Chacoan peccary White-lipped peccary Collared peccary

20 26 30

Benirschke and Kumamoto (1989) Benirschke and Kumamoto (1989) Benirschke and Kumamoto (1989)

572 572 573

Hippopotamus

36

Stanyon (Unpublished)

573

Bactrian camel/Two-humped camel Dromedary camel/One-humped camel

74 74

Graphodatsky (Unpublished) CRES San Diego (Unpublished)

574 575

Lesser malay chevrotain/Asiatic mouse deer

32

Gallagher et al. (1996)

576

45–46 30

CRES San Diego (Unpublished) CRES San Diego (Unpublished)

577 577

58

CRES San Diego (Unpublished)

578

46–48 6F/7M 14M 46 46 8F 8F/9M 68 68 68 56 66 66 66 68 68 68 70 70 + B’s

CRES San Diego (Unpublished) CRES San Diego (Unpublished) Fu and Yang (Unpublished) Yang (Unpublished) Yang et al. (1995) Fu and Yang (Unpublished) Fu and Yang (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) Graphodatsky (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) Graphodatsky (Unpublished) Graphodatsky (Unpublished) CRES San Diego (Unpublished) Graphodatsky (Unpublished)

579 579 580 580 581 582 582 583 584 585 586 587 588 589 590 591 592 593 594

58

CRES San Diego (Unpublished)

595

55 32 38 34F/33M 32F/31M 32F/31M 46 50

CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) Gallagher et al. (1998) Iannuzzi (1994)

596 597 598 599 599 600 601 602

Okapi Masai giraffe Siberian musk deer

Family Cervidae Elaphodus cephalophus Muntiacus muntjak vaginalis Muntiacus feae Muntiacus reevesi Muntiacus reevesi Ideogram Muntiacus gongshanensis Muntiacus crinifrons Dama mesopotamica (dama) Axis kuhlii Axis calamianensis Cervus unicolor malaccensis Cervus nippon hortulorum Cervus nippon mandarinus Cervus albirostris Cervus elaphus bactrianus Cervus elaphus sibiricus Alces alces Rangifer tarandus Capreolus pygargus

Tufted deer North Indian muntjac/Barking deer Fea’s muntjac/Barking deer Chinese muntjac/Barking deer Chinese muntjac/Barking deer Gongshan muntjac/Barking deer Black muntjac/Barking deer Persian fallow deer Kuhl’s deer/Axis deer Calamian deer/Axis deer Malayan sambar deer/Wapiti Dybowski’s sika Mandarin sika White-lipped deer/Thorold’s deer Bactrian wapiti/Red deer Red deer/Wapiti Moose/Elk Siberian reindeer/Caribou Roe deer

Family Antilocapridae Antilocapra americana

Pronghorn

Family Bovidae Tragelaphus angasii Tragelaphus strepsiceros Tragelaphus imberbis Tragelaphus eurycerus Taurotragus oryx pattersonianus Taurotragus derbianus Boselaphus tragocamelus Bubalus bubalis

Lowland nyala Greater kudu Lesser kudu Bongo Patterson’s eland Giant eland Nilgai Asian water buffalo

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

xxvi

CONTENTS

SPECIES

COMMON NAME

Bubalus bubalis Ideogram Bubalus depressicornis Syncerus caffer Bos taurus

Asian water buffalo Lowland anoa Forest buffalo/African buffalo Domestic cattle

Bos taurus Ideogram Bos javanicus Bos frontalis (gaurus) Bos grunniens

Domestic cattle Javan banteng/Bali cattle Gaur/Seladang Yak/Oxen

Bison bonasus Cephalophus rufilatus Cephalophus niger Cephalophus sylvicultor Cephalophus dorsalis Cephalophus monticola Kobus ellipsiprymnus defassa Kobus ellipsiprymnus ellipsiprymnus Kobus megaceros Kobus leche Kobus kob thomasi Pelea capreolus Hippotragus equinus Hippotragus niger Oryx dammah Oryx leucoryx Oryx gazella beisa Oryx gazella callotis Oryx gazella gazella Addax nasomaculatus Damaliscus hunteri Damaliscus pygargus phillipsi Damaliscus lunatus topi Alcelaphus buselaphus jacksoni Connochaetes gnou Connochaetes taurinus albojubatus Oreotragus oreotragus stevensoni Raphicerus campestris Neotragus moschatus Madoqua guentheri Madoqua kirkii Antilope cervicapra Aepyceros melampus petersi Gazella saudiya Gazella bennettii Gazella gazella Gazella spekei Gazella cuvieri Gazella rufifrons laevipes Gazella subgutturosa subgutturosa

European bison/Wisent Red-flanked duiker Black duiker Yellow-backed duiker Bay duiker Blue duiker Defassa waterbuck Ellipsen waterbuck Nile lechwe Red lechwe Uganda kob Rhebok Roan antelope Sable antelope Scimitar-horned oryx Arabian oryx Beisa oryx Fringe-eared oryx Gemsbok Addax Hunter’s hartebeest Blesbok/Sassabie Topi Jackson’s hartebeest White-tailed gnu/Black wildebeest Brindled gnu Klipspringer Steenbok Suni antelope/Dwarf antelope Guenther’s dik-dik Kirk’s dik-dik Blackbuck Black-faced impala Saudi gazelle Chinkara/Gazelle/Indian gazelle Mountain gazelle Speke’s gazelle Cuvier’s gazelle Red-fronted gazelle Persian goitered gazelle

2N

SOURCE*

50 48 54–56 60

Iannuzzi (1994) CRES San Diego (Unpublished) CRES San Diego (Unpublished) Graphodatsky and Biltueva (Unpublished) Cribiu et al. (2001) CRES San Diego (Unpublished) CRES San Diego (Unpublished) Graphodatsky and Biltueva (Unpublished) Graphodatsky (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) Kingswood et al. (1998a) Kingswood et al. (1998a) Kingswood et al. (2000) Kingswood et al. (2000) Kingswood et al. (2000) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) Kumamoto et al. (1999) CRES San Diego (Unpublished) Kumamoto et al. (1999) Kumamoto et al. (1999) CRES San Diego (Unpublished) CRES San Diego (Unpublished) Kumamoto et al. (1996) Kumamoto et al. (1996) Kumamoto et al. (1996) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) Kingswood et al. (1998b) Kingswood and Kumamoto (1996) Kingswood and Kumamoto (1997) CRES San Diego (Unpublished) CRES San Diego (Unpublished) Kumamoto et al. (1995) Kumamoto et al. (1995) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished)

60 60 58 60 60 60 60 60 60 60 53–54 50–52 52 48 50 56 60 60 56–58 57–58 58 58 56 58 44 38 36 40 58 58 60 30 52–56 48–50 46–48 30–33 58–60 46–53 49–52 34F/35M 32F/33M 32F/33M 58 32F/31M

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

603 604 605 606 607 608 609 610 611 612 613 614 615 616 617 617 618 619 620 621 622 623 624 625 626 627 628 629 630 631 632 633 634 634 635 636 636 637 638 639 640 641 641 642 642 643 643 644

CONTENTS

SPECIES

COMMON NAME

2N

Gazella subgutturosa marica

Sand gazelle

30–32F 31–33M 58 30F/31M 38–40 38–40 34–39 56 42 50 56 52 48

Gazella thomsonii Gazella granti roosevelti Gazella dama ruficollis Gazella dama mhorr Gazella soemmerringii Antidorcas marsupialis Oreamnos americanus Capricornis crispus Naemorhedus goral Budorcas taxicolor Ovibos moschatus

Thomson’s gazelle Roosevelt’s gazelle Addra gazelle Mhorr gazella Soemmerring’s gazelle Springbok/Springbuck Rocky mountain goat Serow Goral Takin Muskox

Capra hircus

Domestic goat

60

Capra cylindricornis Capra pyrenaica hispanica Pseudois nayaur szechuanensis Ovis orientalis musimon Ovis aries Ovis aries Ideogram Ovis canadensis

East Caucasian tur/Goat Spanish ibex/Goat Chinese bharal/Blue Sheep European mouflon/Sheep Domestic sheep Domestic sheep Rocky mountain bighorn sheep

60 60 56 54 54 54 54

SOURCE* CRES San Diego (Unpublished)

644

CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) Biltueva et al. (Unpublished) Biltueva et al. (Unpublished) CRES San Diego (Unpublished) Biltueva and Graphodatsky (Unpublished) Biltueva and Graphodatsky (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) Cribiu et al. (2001) Cribiu et al. (2001) CRES San Diego (Unpublished)

645 645 646 646 647 648 648 649 650 651 652

Diceros bicornis

653 654 655 656 657 658 659 660 661

Order Perissodactyla

Family Equidae Equus asinus Equus asinus (africanus) somaliensis Equus hemionus onager Equus hemionus kulan Equus przewalskii Equus caballus Equus caballus Ideogram Equus burchelli Equus quagga boehmi Family Tapiridae Tapirus terrestris Tapirus pinchaque Tapirus bairdii Tapirus indicus Family Rhinocerotidae Rhinoceros unicornis Ceratotherium simum simum

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Donkey Somali wild ass Onager Kulan Przewalski’s wild horse Domestic horse Domestic horse Burchell’s zebra Grant’s zebra/Quagga

62 62–64 55–56 54–55 66 64 64 44 44–45

Alaoui et al. (2004) Houck et al. (1998) CRES San Diego (Unpublished) CRES San Diego (Unpublished) CRES San Diego (Unpublished) Stanyon (Unpublished) Bowling et al. (1997) Graphodatsky (Unpublished) CRES San Diego (Unpublished)

665 665 666 667 668 669 670 671 672

Lowland tapir Mountain tapir Baird’s tapir Malayan tapir

80 76 80 52

Houck et al. (2000) Houck et al. (2000) Houck et al. (2000) Houck et al. (2000)

673 674 675 676

Indian rhinoceros Southern white rhinoceros/Square-lipped rhinoceros Black rhinoceros

82 82

Houck et al. (1994) Houck et al. (1994)

677 678

84

Houck et al. (1995)

679

*Source: Refer to References for published karyotypes and to Contributors for unpublished karyotypes

FOREWORD I

n this “Era of the Genome,” when we obtain ever greater insight into our own genetic makeup and that of other species, the publication of a new Atlas of Mammalian Chromosomes may seem to be truly anachronistic. There are some of us, however, who actually believe not only that the genes are important but also that many aspects of a fairly good evolutionary heritage of extant mammals may be derived from their chromosomal structure, from their number and their centromere position, as well as from their banding details. Thus, this new effort by Stephen J. O’Brien, Joan C. Menninger, and William G. Nash is one that was undertaken with much devotion and enormous perseverance. It updates all of the information from hundreds of publications that are so difficult to access. Many years ago now, the late T. C. Hsu and I endeavored to bring together whatever we could obtain about karyotypes in 10 volumes on mammalian chromosome structure. It was published as an atlas by Springer-Verlag in New York HSU and Benirschke (1971; 1973; 1974; 1975; 1977). For the presentation in that atlas, however, banding cytological techniques had not been well developed, so that only in the last few years was it possible to include some chromosome-banding pictures; those studies were then just evolving. But the task of assembling annually dozens of new mammalian chromosome structures became quickly so overwhelming that we ceased the publication in 1977. However, the origin of that initial effort is perhaps of historical interest. In the 1960s, the annual meetings of FASEB were held in Atlantic City. This was a time when one met old friends on the Boardwalk, when new scientific acquaintances were made, and, importantly, when one was able to view the newest books and instruments in the fabulous exhibit hall of the convention center. T. C. Hsu and I met there on many occasions. Once we discussed how best to publish all the new chromosomal information that we were accumulating because nobody wanted to print information on single species forever. Because I had known the late Dr. Heinz Götze of Springer-Verlag for a long time, we were able to persuade him and his company to produce an annual loose-leaf Atlas of Mammalian Chromosomes (1967–1977). So, with the help of many zoos and many collectors and scientists from many regions, these old pages were produced. They have been useful to an extent, but they also really needed radical improvements and, especially, more detailed analysis of banding structures have become necessary. C-bands and G-bands and later R-bands were being developed and new insights were gained in our understanding of the possible relationships of chromosome

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FOREWORD number and structure among related taxa. How did new species evolve, and how did their chromosomal arrangements change? Having been a Robertsonian fusion believer all my life, the information gathered then favored that mechanism as a major evolutionary force—at least so I think. Fission, while doubtless occurring occasionally, has been found to be exceedingly rare in human chromosomal studies, clearly the most frequently analyzed cells. But clear-cut decisions of the chromosomal evolution of a variety of mammals is still to come; chromosome painting and fluorescence in situ hybridization (FISH) studies are now helpful in many cases, as have been the much improved banding details. Despite all of that, the controversy of “fission versus fusion” still exists, and it will interest us for some time to come. One problem for the working scientist remained, though, the need to gather numerous articles for access to this chromosomal information, a laborious process. Thus, I welcome the enormous effort that Stephen J. O’Brien and colleagues have made in producing this new atlas for us. O’Brien, of course, is well known to geneticists as the editor of the Journal of Heredity. Others will know him because of his seminal contributions on the evolution of mammalian viruses and the adaptation to this exposure by the host. Still others know Steve as the author of his recent book, Tears of the Cheetah (2003), in which he recalls his growth as a genetic scientist and, especially, the discovery of the homogeneity of the genetic make-up of cheetahs. Steve has also been instrumental in participating in the creation of the new taxonomy of Mammalia. This aspect is heavily relied upon in this large new volume before you. It will also perhaps be confusing to older generations of scientists. But the new atlas makes the need for this new arrangement easy to comprehend and to accept it as an important principle of taxonomy. The chromosomes the editors were able to gather for this comprehensive atlas come from their own studies and those of numerous colleagues, as were the ones we prepared in the former atlas. Thus, there are some differences in the quality of the preparations, but all of their structural features are easily grasped, and there are a sufficient number of references that allow one to go back to the original articles, if one is so disposed. And, of course, when one has a really overriding interest, for most taxa there are fibroblast cultures available from the authors. It becomes thus easily possible to proceed with additional karyological or genetic studies. This atlas is a welcomed collection of most of the information on what is currently known of mammalian karyotypes, and it will be widely used as a reference volume. It must have been a monumental task to bring this much material together, and we thus congratulate the editors and hope that these pages will find a wide distribution among scientists. KURT BENIRSCHKE San Diego, California September 2005

PREFACE U

nderstanding the history of life on Earth has become invigorated by technological advances in molecular evolution, in fossil inference, and in geology and by comparative descriptions of anatomy, physiology, and behavior. The earliest mammals were the morganucodontids, mouse-sized insectivores that scurried around in the shadows of dinosaurs across the Triassic and Jurassic epochs. With articulated jaws and squared forelimbs adapted for running, the tiny shrewish creatures never got much bigger than a cat since their first appearance in the fossil record 210 million years ago (mya) until an asteroid or comet blackened Earth 65 mya and eliminated the giant dinosaurs forever. By the early Eocene (55 mya), recognizable adaptations to the world ecological niches were appearing in precursors to the 26 orders of modern mammals. Recently, sophisticated phylogenetic analyses of large deoxyribonucleic acid (DNA) sequence datasets across mammalian species, interpreted in the context of paleontological remains and tectonic movements, have revealed an evolutionary hierarchy we call the mammalian radiations (see end papers). The timing and rationale for taxon divergence, for species isolation, and for adaptation remain imprecise, but genomic tools have already exceeded modest expectations in revealing secrets of our ancestors. Today, around 4700 species of mammals survive on Earth. They range from the smallest 1.5-g Kitti’s hog-nosed bat to the 190,000-kg blue whale, the largest species to ever live. They reflect as diverse an assemblage of anatomic, behavioral, and physiological adaptation as the world has ever seen. The assembly instructions, retained for each species by natural selection across a 200-million-year history, are preserved in modern genomes. The field of comparative genomics is providing a new opportunity to annotate the distinctions, divergence, and genetic occurrences that have sculptured modern genomes. (O’Brien et al., 1999; Murphy et al., 2005) The rich diversity of living mammals is aptly chronicled in several outstanding monographs that describe adaptive specialization and taxonomic classification (Ewer, 1973; Eisenberg, 1981; Novak, 1999; Wilson and Reeder, 1993; 2005; MacDonald, 2001). Further, the paleontological literature reveals the anatomic transitions that characterize each lineage of mammal evolution (McKenna and Bell, 1997; Savage and Russell, 1993). The genetic scripts for each of evolution’s creations are encoded in the chromosomes of living species, organized in genomes containing nearly 3 billion nucleotide letters as instructions to make a manatee, a naked mole rat, a vampire bat, a lion, an elephant, or a cognitive human being. We present in this atlas a sampling of 815 species

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PREFACE karyotypes, most (but not all) G-banded, providing a view of chromosomal bar codes of homologous segments among related species. The karyotype photographs provide a starting point for genomic inquiry, a platform for genetic discovery. This Atlas of Mammalian Chromosomes is a sequel to the pioneering volumes edited by T. C. Hsu (now deceased) and Kurt Benirschke (author of the foreword). In 10 editions, unbanded karyotypes for hundreds of species were archived. Here we extend that beginning with the best karyotypes available for 815 species. We asked experts on cytogenetics for each order of mammals to compose a short overview on the biology, genetics, cytogenetic accomplishments, and analyses for each order. For each species we present the best karyotype available, the common and Latin name for the species, the published citation, and contributing author(s). The collection offers in one place an unabridged description of mammalian cytogenetics from a half century of work by hundreds of cytogenetic artisans. The Human Genome Project, an international collaboration to determine the full DNA sequence of a human genome, published a near-finished version in October 2004 (International Human Genome Sequencing Consortium, 2004). The dense genome-mapping annotation and redundant sequencing immediately elevated the human genome as the primary mammal for genomic studies. The new human sequence showed our genomes to be 2.85 billion base pairs in length, around 2% protein coding, around 50% repetitive elements, and containing 20,000–25,000 protein-coding genes. Full genome sequences of other mammalian species recently sequenced (mouse, rat, bovine, dog) showed that other mammals had very similar composition statistics and gene order, affirming the indisputable unity of all living mammals. Our genomes all descend from an ancient 200-million-year-old ancestor, the tiny insectivore of the Triassic. Comparative genomics of mammals is a fast-moving new field empowered by the vastness of genome information, by powerful computer routines developed to quantify DNA divergence, and by a potential to unlock the hidden secrets in the human and related species genomes. Comparative genomics seeks two broad goals: annotation of the mysterious operatives of the human genome and unraveling the changes and adaptations that led to mammalian species diversification. These are heady issues and only very recently have geneticists actually had the opportunity, data, and tools to pursue such profound genetic challenges. Chromosome identification is a century old, while G-banding and homology segment identification first appeared in the 1970s. Chromosome painting (also called Zoo-FISH, for fluorescent in situ hybridization) has allowed the characterization of homologous chromosome segments virtually by direct observation. Over 50 species of mammals have been “painted” with flow-sorted human and other mammalian chromosome libraries as the discipline of phylogenomics (the study of chromosome exchanges that characterize mammalian ordinal lineages) is beginning to bear fruit (O’Brien et al., 1999; VandeBerg and Marshall Graves, 1999). The ultimate comparative genomics tool, an assembled whole genome sequence, has been proposed and developed for 10 species of mammals: human, chimpanzee, orangutan, macaque, mouse, rat, cow, dog, opossum, and platypus (www.nhgri.nih.gov). Whole-genome sequence coverage has been funded for eight species in order to capture the evolutionary diversity of eight additional mammal species: African savanna elephant, lesser hedgehog tenrec, nine-banded armadillo, rabbit, domestic cat, hedgehog, guinea pig, and European common shrew. Before long a new mammal genome will

PREFACE appear every month or even each week. A deluge of genomic sequence data has geneticists of all specialties anticipating the assembly, comparison, analysis, and mining of mammalian genomes. The atlas presented here is an early salvo in the compiling of genomic information. Our understanding of the details of species formation of developmental specialization, of the exact targets of natural selection, of life’s darkest secrets is one step closer than before. STEPHEN J. O’BRIEN JOAN C. MENNINGER WILLIAM G. NASH Frederick, Maryland February 2006

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ACKNOWLEDGMENTS W

e wish to thank all those contributors (p. xxxvii) who donated their time, effort, and encouragement to this major collection of mammalian karyotypes. In particular we are indebted to all who cheerfully who sent reprints, karyotypes, electronic images, text, and other information incorporated in this edition. Without all the technical help and support from the collaborators and authors, this work would not have reached fruition.

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CONTRIBUTORS DAVID ADLER Department of Pathology University of Washington Seattle, Washington V. M. ANISKIN Severtsov Institute of Ecology and Evolution Russian Academy of Sciences Moscow, Russia

VIOLETTE BEKLEMESHEVA Siberian Department Institute of Cytology and Genetics Russian Academy of Sciences Novosibirsk Russia KURT BENIRSCHKE Department of Pathology UCSD Medical Center San Diego, California

ULFUR ARNASON University of Lund Division of Evolutionary Molecular Systematics Lund, Sweden

JOHN W. BICKHAM Department of Wildlife and Fisheries Sciences Texas A&M University College Station, Texas

ROBERT J. BAKER Department of Biological Sciences Texas Tech University Lubbock, Texas

FRANCECCA BIGONI Genetics Branch, Center for Cancer Research National Cancer Institute Frederick, Maryland

KRISTEN BALCH Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California

LARISA BILTUEVA Siberian Department Institute of Cytology and Genetics Russian Academy of Sciences Novosibirsk Russia

IRINA BAKLUSHINSKAYA Siberian Department Institute of Cytology and Genetics Russian Academy of Sciences Novosibirsk Russia

GARY BRONNER Department of Mammals Transvaal Museum Pretoria, South Africa

C. M. BARROSO Departamento de Genetica Universidade Federal do Para Braszl

D. L. BUSBEE Department of Anatomy and Public Health College of Veterinary Medicine Texas A & M University College Station, Texas

BETH BAUM Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California

SUELLLEN CHARTER Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California

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CONTRIBUTORS

LEONA CHEMNICK Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California RAMESH C. CHOUDHURY Cytogenetics and Genetic Toxicology Laboratory Department of Zoology Berhampur University Berhampur Orissa, India RENEE COOK Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California CRES Center for Reproduction of Endangered Species Zoological Society of San Diego San Diego, California CHEN DAOQUAN Institute of Hydrobiology Academia Sinica Wuhan, Hubei, China

UTA FRANCKE Department of Genetics Stanford School of Medicine Stanford, California LUTZ FRÖENICKE Population Health and Reproduction School of Veterinary Medicine University of California, Davis Davis, California JULIE FRONCZEK Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California BEIYUAN FU Kunming Institute of Zoology The Chinese Academy of Science Yunnan, China DONALD GALLAGHER Department of Pathology Pfizer, Inc. Groton, Connecticut MILTON H. GALLARDO Instituto de Ecologia y Evolucion Universidad Austral de Chile Valdivia, Chile

G. DOBIGNY Laboratoire Origine Structure et Evolution de la Biodiversite Museum National d’ Histoire Naturalle Paris, France

WILMA GEORGE Lady Margaret Hall Department of Zoology Oxford, United Kingdom

MARY ANN DOMINGOS Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California

ALEXANDER S. GRAPHODATSKY Siberian Department Institute of Cytology and Genetics Russian Academy of Sciences Novosibirsk Russia

MARK D. B. ELDRIDGE Department of Biological Sciences Macquarie University Sydney, Australia and Evolutionary Biology Unit Australian Museum Sydney, Australia

DARIA GRAPHODALSKAYO ETN Zurich, Switzerland

MALCOLM A. FERGUSON-SMITH Centre for Veterinary Science University of Cambridge Cambridge, United Kingdom

I. F. GREENBAUM Department of Biology Texas A&M University College Station, Texas

I. FERRARI Departamento de Genetica Faculdade deMedicina de Ribeirao Preto Universidade de Sao Paulo Sao Paulo, Brazil

BRIAN A. GRAY Department of Pediatrics Division of Genetics University of Florida Gainesville, Florida

CONTRIBUTORS FRANK GRUˇ TZNER Comparative Genomics Research Group Research School of Biological Sciences Australian National University Canberra, Australia

STEVEN KINGSWOOD Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California

SUSAN HANSEN Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California

ARLENE KUMAMOTO (DECEASED) Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California

H. HAYES Laboratoire de Génétique Biochimique et Cytogenetique Jouy-en Josas, France

TANGLIANG LI Centre for Veterinary Science University of Cambridge Cambridge United Kingdom and Kunming Institute of Zoology Chinese Academy of Science Yunnan, China

D. L. HAYMAN Department of Zoology and Genetics University of Adelaide Adelaide, Australia KRISTOFER M. HELGEN School of Earth and Environmental Sciences University of Adelaide Adelaide, Australia MARLYS L. HOUCK Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California LEOPOPLDO IANNUZZI National ResearchCouncil (CNR)/IABBAM Naples-Ponticelli, Italy ANDREA JOHNSON Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California P. G. JOHNSTON School of Biological Sciences Macquarie University North Ryde, Australia WILLIAM JORGE Departmento de Biologia Geral Instituto de Ciencias Biologicas Universidade Federal de Minas Gerais (ICB-UFMG) Caixa 486, 20.161–970 Belo Horizonte, Minas SHIN-ICHIRO KAWADA School of Agricultural Sciences Nagoya University Nagoya, Japan

ELENA LYAPUNOVA Siberian Department Institute of Cytology and Genetics Russian Academy of Sciences Novosibirsk Russia M. MACHOLAN Institute of Systematic and Ecological Biology Academy of Sciences Brno, Czech Republic GRACE MAGEE Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California JENNIFER A. MARSHALL GRAVES Comparative Genomics Research Group Research School of Biological Sciences Australian National University Canberra, Australia GRESSA MCDOWELL Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California G. M. MCKAY School of Biological Science Macquarie University North Ryde, Australia

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CONTRIBUTORS LEON R. MCQUADE Stem Cell Group Diabetes Transplant Unit Prince of Wales Hospital Randwick, Australia JOAN C. MENNINGER Laboratory of Genomic Diversity National Cancer Institute SAIC-Frederick Frederick, Maryland C. J. METCALFE School of Molecular and Cell Biology University of Connecticut Storrs, Connecticut ANDREA MIKOLON Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California Macquarie University North Ryde, Australia D. A. MILLER Department of Molecular Biology and Genetics Wayne State University School of Medicine Detroit, Michigan CHEN MINRONG Institute of Hydrobiology Academia Sinica Wuhan, Hubei, China WILLIAM MODI Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California E. MORIELLE-VERSUTE Institute of Biosciences UNESP Sao Jose do Rio Preto, Sao Paulo, Brazil J. D. MURRAY School of Biological Science Macquarie University North Ryde, New South Wales, Australia C. Y. NAGAMACHI Departamento de Genetica CCB Universidade Federal do Para Campus do Guama Belem, Brazil

WILLIAM G. NASH H & W Cytogenetic Services, Inc. Lovettsville, Virginia and Laboratory of Genomic Diversity National Cancer Institute-Frederick Frederick, Maryland T. B. NESTEROVA Siberian Department Institute of Cytology and Genetics Russian Academy of Sciences Novosibirsk, Russia EVIATAR NEVO University of Haifa Mount Carmel, Haifa, Israel WENHUI NIE Laboratory of Cellular and Molecular Evolution Kunming Institute of Zoology Chinese Academy of Sciences Kunming, Yunnan, China STEPHEN J. O’BRIEN Laboratory of Genomic Diversity National Cancer Institute Frederick, Maryland SEN-ICHI ODA School of Agricultural Sciences Nagoya University Nagoya, Japan HESED PADILLA-NASH Genetics Branch National Cancer Institute National Institutes of Health Bethesda, Maryland S. PATHAK Department of Biology University of Texas System Cancer Center M.D. Anderson Hospital and Tumor Institute Houston Texas JELINDA PEPPER Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California POLINA PERELMAN Laboratory of Genomic Diversity National Cancer Institute Frederick, Maryland

CONTRIBUTORS ABEL PONCE DE LEON Department of animal science College of Agriculture Food & Environmental Science University of Minnesota St. Paul, Minnesota

MARGARITA ROGATCHEVA Siberian Department Institute of Cytology and Genetics Russian Academy of Sciences Novosibirsk Russia

MONTSERRAT PONSA Department Biologia Cellular Universitat Autonoma de Barcelona Cerdanyola del Valles, Spain

NIKOLAI B. RUBTSOVA Siberian Department Institute of Cytology and Genetics Russian Academy of Sciences Novosibirsk Russia

PATRICIA A. POORMAN-ALLEN Department of Anatomy Duke University Medical Center Durham, North Carolina

YVES RUMPLER Université Louis-Pasteur Institut d’Embryologie, Faculté de Médecine Strasbourg Cedex, France

NICHOLAS C. POPESCU Molecular Cytogenetics Section Laboratory of Experimental Carcinogenesis National Cancer Institute Bethesda, Maryland

OLIVER A. RYDER Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California

MAZIN B. QUMSIYEH Genetics Department Yale University School of Medicine New Haven, Connecticut

OLGA SABLINA Siberian Department Institute of Cytology and Genetics Russian Academy of Sciences Novosibirsk Russia

SEVIL RADJABLI Siberian Department Institute of Cytology and Genetics Russian Academy of Sciences Novosibirsk Russia WILLEM RENS Centre for Veterinary Science University of Cambridge Cambridge, United Kingdom ERIC A. RICKART Utah Museum of Natural History University of Utah Salt Lake City, Utah TERENCE J. ROBINSON Evolutionary Genomics Group Department of Zoology University of Stellenbosch Matieland, South Africa ALFRED L. ROCA Laboratory of Genomic Diversity National Cancer Institute SAIC-Frederick Frederick, Maryland RUTH ROFE LeCornu Street South Australia, Australia

HITOSHI SATOH Division of Pathology Department of Cancer Biology Institute of Medical Science University of Tokyo Tokyo, Japan O. L. SEROV Siberian Department Institute of Cytology and Genetics Russian Academy of Sciences Novosibirsk Russia HECTOR N. SEUANEZ Genetics Section Instituto Nacional de Cancer Rio de Janeiro, Brazil H. SOMA Department of Obstetrics and Gynecology Tokyo Medical College Tokyo, Japan A. E. SPOTORNO Laboratorio de Citogenetica Evolutiva Departamento de Biologia Celular y Genetica Facultad de Medicina Universidad de Chile Santiago, Chile

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CONTRIBUTORS K. S. SREEPADA Department of Applied Zoology Mangalore University Mangalagangothri Mangalore, Karnataka India ROSCOE STANYON Center for Cancer Research National Cancer Institute Frederick, Maryland DEAN A. STOCK Department of Biology University of Texas System Houston, Texas GERALD STRANZINGA ETN Zurich, Switzerland MARTA SVARTMAN Comparative Molecular Cytogenetics Core Genetics Branch National Cancer Institute-Frederick Frederick, Maryland EMMA TEELING Laboratory of Genomic Diversity National Cancer Institute-Frederick Frederick, Maryland JR., CHARLES S. THAELER Department of Biology New Mexico University Las Cruces, New Mexico MARIANNE VOLLETH Department of Human Genetics University of Erlangen Erlangen, Germany V. T. VOLOBOUEV Laboaratoire de Zoologie Mammiferes et oiseaux Museum National d’Histoire Naturelle Paris, France JACLYN M. WATSON LaTrobe University Bundoora, Victoria, Australia ROBERT WAYNE Department of Organismic Biology Ecology and Evolution University of California Los Angeles, California

JOHANNES WIENBERG Institute of Human Genetics Ludwig Maximilian University Munich, Germany DON WILSON Division of Mammals National Museum of Natural History Smithsonian Institution Washington, D.C. KRISTY WOLFE Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California JAMES WOMACK Department of Veterinary Pathology Texas A&M University College Station, Texas TAMMY WRIGHT Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego San Diego, California DORIS H. WURSTER-HILL Department of Pathology Dartmouth Medical School Hanover, New Hampshire FENTANG YANG Centre for Veterinary Science University of Cambridge Cambridge, United Kingdom and Kunming Institute of Zoology Chinese Academy of Science Yunnan, China JORGE YUNIS Jefferson Medical School Department of Pathology Philadelphia, Pennsylvania GUAN ZHIMERI Institute of Hydrobiology Academia Sinica Wuhan, Hubei, China

MONOTREMATA Order Monotremata Chromsomes of Monotreme Mammals Monotremes (mammalian subclass Prototheria) are usually described as the earliest offshoot of the mammalian lineage, having diverged about 210 million years ago (mya) from therian mammals (eutherians and marsupials) (Woodbourne et al., 2003; reviewed by Grützner and Graves, 2004; Musser, 2003). Generally classified in a single order Monotremata of the mammalian subclass Prototheria, only three extant species are known: the duck-billed platypus (Ornithorhynchus anatinus) and two echidna species, the Australian short-beaked echidna (Tachyglossus aculeatus) and the long-beaked Niugini echidna (Zaglossus bruijni), which has recently been split into three different species on the basis of morphological diversity (Flannery and Groves, 1998). The fossil record of monotremes in Australia, along with a single tooth from South America, shows that monotremes, in a variety of forms, were widespread in eastern Gondwana (Musser, 2003; Pascual et al., 1992). Molecular comparisons suggest that the two extant echidna species are closely related (~3 million years since divergence) but that echidnas are distantly related to the platypus (estimates from 20 to 45 million years; Cao et al., 1998; Kirsch and Mayer, 1998). Monotremes are confined to Australia and New Guinea. The platypus inhabits freshwater streams, lakes, and lagoons of eastern Australia, in Queensland, New South Wales, Victoria, and Tasmania. The short-beaked echidna is found in all major habitats in mainland Australia and Tasmania. Both species are relatively common, although the platypus is considered to be vulnerable because of riverbank erosion and water quality. The long beaked echidna occurs only in New Guinea, where it is highly threatened by habitat destruction and other human factors. Monotremes feature a unique mixture of mammalian, avian, reptilian, and specialized morphological and physiological features. Because of their basal evolutionary position among mammals, they are invaluable for comparative genomics. The study of monotreme chromosomes and genes has provided many insights into mammalian sex chromosome evolution and the origin of the mammalian genome (Grützner et al., 2003; Grützner et al., 2004; Grützner and Graves, 2004). Early cytologists considered the karyotypes of the platypus and echidna to be similar to those of reptiles and birds, which contain tiny dotlike

Atlas of Mammalian Chromosomes, Edited by Stephen J. O’Brien, Joan C. Menninger, William G. Nash Copyright © 2006 John Wiley & Sons, Inc.

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MONOTREMATA microchromosomes as well as normal-sized macroelements (Matthey, 1949; White, 1973). However, Van Brink (1959) disagreed, noting that the small chromosomes of monotremes were much larger than sauropsid microchromosomes and there was a more continuous distribution of chromosome size than the bimodal distribution in birds and reptiles. No consistent chromosome numbers were available until Bick and Jackson (1967) described the karyotypes of the platypus and echidna. They correctly reported that the echidna has 2n = 63 in males and 2n = 64 in females but reported incorrect numbers for the platypus, later corrected to 2n = 52 in both sexes (Bick and Sharman, 1975). The chromosomes of the platypus and echidna were described in greater detail by Murtagh (1977), and G-band and R-band karyotypes and accounts of replication banding and activity were published by Wrigley and Graves (1988a,b). The discovery of a multivalent chain in male meiosis of monotremes was consistent with the curious observation that several chromosomes in the monotreme karyotype seemed to lack an exact homologue. Platypus males were described as having a chain of eight chromosomes and echidna a chain of nine at meiosis (Bick, 1992; Murtagh, 1977). The meiotic chain in monotremes provides the only naturally occurring example of translocation chains in a vertebrate genome, though such systems are known in several plants and invertebrates. Interpretation of the monotreme meiotic chain was therefore based on analogy to multivalent meiotic chains described in classic work on the evening primrose Oenothera (Cleland, 1962), in which meiotic chains or rings result from balanced heterozygosity for autosomal translocations. The chain is formed when homologous regions of each translocated chromosome pair with two different partners, and recombination occurs in the homologous regions at the terminal part of the chromosomes. In insect and crustacean groups, translocation heterozygosity involves the sex chromosomes (Syren and Luykx, 1977; Luykx and Syren, 1981). The only chain element that could be identified with traditional cytological methods was an X chromosome, defined by its presence in two copies in males and a single copy in females. Comparative mapping showed that this chromosome shared homology with part of the human X chromosome (Watson et al., 1990, 1991). This X chromosome shows asynchronous deoxyribonucleic acid (DNA) replication suggestive of X chromosome inactivation (Wrigley and Graves, 1988b), and dosage compensation has been confirmed by measuring transcription rates (Grützner et al., 2003). The monotreme chain was considered to represent a complex sex chromosome system. How this system worked was challenging to discover because the difficulty of identifying the smaller chromosomes and the striking chromosome heteromorphism made male–female comparisons nearly impossible by classical cytology. It was concluded from comparisons of male and female karyotypes in both species that males and females shared the same set of small unpaired chromosomes (Wrigley and Graves, 1988a). More recently all platypus chromosomes were isolated by flow sorting and tested by chromosome painting on male and female metaphase chromosomes (Rens et al., 2004) as well as on meiotic chains and sperm (Grützner et al., 2004). This revealed that the platypus has a X1Y1X2Y2X3Y3X4Y4X5Y5 male–X1X1X2X2X3X3X4X4X5X5 female sex chromosome system. In males these 10 sex chromosomes adopt an XY alternating pattern and segregate to 5X (female-determining) and 5Y (male-determining) sperm. It is likely that the tiny Y5 has been lost in the related echidna, which has a meiotic chain of nine chromosomes. However, the difference in the number of chromosomes in the platypus and echidna indicates that numerous as-yet

ORDER MONOTREMATA unidentified chromosome rearrangements have occurred to separate these species. In both monotreme species, chromosomes appear in a defined order in the fibrillar sperm head, with the X near the apex (Watson et al., 1996; Graves et al., 2003).

Mitotic Chromosomes of Platypus (Ornithorhynchus anatinus, Order Monotremata of Mammalian Subclass Prototheria) The platypus has 2n = 52 in both sexes (Bick and Sharman, 1975; Murtagh, 1977; Wrigley and Graves, 1988b). The two members of homologous pairs frequently show differences in size and morphology. The largest six autosomes and a large X chromosome are readily identified by size, morphology, and banding patterns. The heterochromatic and heteromorphic short arm of chromosome 6 bears the nucleolus organizer. Most of the chromosomes of the platypus, however, are small and poorly differentiated by G- or R-banding (Wrigley and Graves, 1988a). They have generally been grouped into just two size classes. It has been particularly difficult to characterize sex differences in platypus mitotic cells. A large X chromosome could be recognized by its presence in two copies in females and a single copy in males. Gene mapping establishes homology with at least a part of the long arm and the pericentric region of the human X chromosome (Watson et al., 1990), but not the medial region of the long arm or the distal short arm (Watson et al., 1991). Using chromosome paints, it has at last been possible to identify all chromosomes of the meiotic chain in the platypus and to examine their pairing relationships (Grützner et al., 2004; Rens et al., 2004). Comparisons between the karyotypes of males and females showed that there are five chromosomes that are present in two copies in females and a single copy in males (i.e., five X chromosomes, including the X previously identified). In addition, there are five chromosomes that are present only in males (i.e., five Y chromosomes). Females are therefore homozygous for all X chromosomes and contain no unpaired elements. The 10 chromosomes that are unpaired in males show partial homologies that suggest they were formed by successive translocation and provide a putative order in the chain (Rens et al., 2004). Chromosome painting at male meiosis shows that the five X chromosomes alternate with the five Y chromosomes to form the chain of 10, starting with the largest X1 (previously shown to be partly homologous to the human X) and finishing with a tiny male-specific element. At the other end of the chain X5 contains the DMET1 gene that is a marker for the bird sex chromosomes, suggesting an evolutionary link between the mammal XY and bird ZW sex chromosome systems. These 10 chromosomes adopt an alternating pattern at meiosis so that all male-specific Ys are segregated to male-determining sperm and all X chromosomes are segregated to female-determining sperm (Grützner et al., 2004). The sex chromosome system is therefore represented formally as X1X1 X2X2 X3X3 X4X4 X5X5 female–X1Y1 X2Y2 X3Y3 X4Y4 X5Y5 male.

Mitotic Chromosomes of Short-Beaked Echidna (Tachyglossus aculeatus, Order Monotremata of Mammalian Subclass Prototheria) The short-beaked (Australian) echidna has 2n = 63 in the male and 2n = 64 in the female. The two members of homologous pairs frequently showed differences in size and morphology. Again, the largest six chromosomes and a large

3

4

MONOTREMATA X are readily identifiable by size, morphology, and banding patterns (Murtagh, 1977; Wrigley and Graves, 1988a). These large chromosomes appear to be identical to the largest platypus chromosomes by G- and R-banding and for chromosomes 1, 2, and X also by gene mapping (Watson et al., 1992b). The heterochromatic and heteromorphic short arm of chromosome 3 as well as chromosome 6 bear nucleolar organizer regions. A large X1 chromosome is present in two copies in females and a single copy in males. G-banding and gene mapping establish homology between the echidna X1 and platypus X1 chromosome (Watson et al., 1992b). There is also a small submetacentric chromosome that is present in two copies in the female and one in the male. It was suggested that this chromosome represents an X2 chromosome that is part of an X1X2Y male–X1X1X2X2 system in this species, explaining the difference in chromosome numbers in male and female (Murtagh, 1977; Wrigley and Graves, 1988b). In males a Y chromosome was identified as a large unpaired element, present in male but not female karyotypes. However, the situation is likely to be much more complex, since these three chromosomes are part of a meiotic chain of nine (Murtagh, 1977; Watson et al., 1992a, 1996). The elements of the echidna chain, delineated by hybridization with telomere-specific oligonucleotides, were counted and measured (Watson et al., 1992a) but could not be identified with respect to the mitotic karyotype. We have identified six chromosomes additional to X1 X2 in females and a total of nine chromosomes in males that lack an obvious homologue in males. Whether these unpaired chromosomes occur also in females must await chromosome painting, but by analogy with the platypus, it is likely that apparently unpaired chromosomes in females are the result of chromosome heteromorphism. Comparison of the sizes of the elements in the echidna chain suggests that they are homologous, with the exception of the tiny element at the end of the platypus chain (Bick, 1992; Grützner et al., 2004), which is lacking in the echidna. The sex chromosome system is therefore most likely to be represented formally as X1X1 X2X2 X3X3 X4X4 X5X5 female– X1Y1 X2Y2 X3Y3 X4Y4 X5O male.

Mitotic Chromosomes of Long-Beaked Echidna (Zaglossus bruijni, Order Monotremata of Mammalian Subclass Prototheria) The long-beaked (New Guinea) echidna Z. bruijni has a mitotic karyotype similar or identical to that of the short-beaked echidna T. aculeatus. The chromosome number is 2n = 63 in the male and 2n = 64 in the female (Wrigley and Graves, 1988a; Bick, 1992). The two members of homologous pairs frequently showed differences in size and morphology. The heterochromatic and heteromorphic short arms of chromosomes 3 and 6 bear nucleolus organizer regions. As for T. aculeatus, there is a large X1 and a small X2 chromosome, each present in two copies in females and a single copy in males. G-banding suggests homology of the Z. bruijni and T. aculeatus X1 with the platypus X1 chromosome (Wrigley and Graves, 1988b). Again, there are nine unpaired chromosomes in the male, including one large male-specific element similar to that observed in T. aculeatus. Eight chromosomes in females seem to have no homologues, but, again, this is likely to be the result of heteromorphism. At meiosis in males, the unpaired elements form a translocation chain of 9, which includes the X1, Y1 and X2 chromosomes and the other six unpaired chromosomes (Bick, 1992). It is expected that segregation from the meiotic translocation chain is similar to that in male platypus. The sex chromosome

ORDER MONOTREMATA system is also represented formally as X1X1 X2X2 X3X3 X4X4 X5X5 female– X1Y1 X2Y2 X3Y3 X4Y4 X5O male. Frank Grützner Willem Rens Jaclyn M. Watson Malcolm A. Ferguson-Smith Jennifer A. Marshall Graves

5

6

MONOTREMATA

Tachyglossus aculeatus (Short-Beaked Echidna/Short-Nosed Echidna)

2n=64(female)

2n=63(male)

Unpaired elements

Unpaired elements

Graves, Watson, and Grutzner (unpublished)

ORDER MONOTREMATA

Zaglossus bruijni (Long-Beaked Echidna/Long-Nosed Echidna)

2n=64(female)

2n=63(male)

Unpaired elements

Unpaired elements

Graves, Watson, and Grutzner (unpublished)

7

8

MONOTREMATA

Ornithorhynchus anatinus (Platypus)

2n=52 (female)

2n=52 (male)

Rens et al. (2004) Contributed by F. Grutzner

*21 chromosome pairs and 10 unpaired chromosomes labeled E1–E10. The order of these chromosomes determined by their pairing regions is indicated by vertical bars.

MARSUPIALIA T

HERE ARE 7 ORDERS AND 22 families of living marsupials, all restricted to Australasia and the Americas. Of the ~300 currently recognized species, most (75%) are endemic to Australia, New Guinea, and the islands of eastern Indonesia. The remaining ~75 species are found in South and Central America, with only a single species occurring in North America. Marsupials were among the first mammals studied cytogenetically, with the earliest reliable chromosome numbers reported in 1922 for an American species and 1923 for several Australasian species. Since then karyotypic data have been published for over 60% of known species, including representatives of all orders and most (91%) families.

Marsupial chromosome evolution shows two dominant and yet contrasting patterns—extreme conservation in some lineages and extensive rearrangement in others (O’Brien et al., 1999). For example, a morphologically similar 2n = 14 karyotype is found in representatives of six of the seven extant marsupial orders. G-banding and cross-species chromosome painting have confirmed the remarkable homology of this 2n = 14 complement across five of these orders (Dasyuromorphia, Didelphimorphia, Diprotodontia, Microbiotheria, Peramelemorphia), including its almost complete conservation for ~70 million years in eight marsupial families (Dasyuridae, Peramelidae, Vombatidae, Burramyidae, Acrobatidae, Marmosidae, Caluromyidae, Microbiotheriidae). The presence of this extraordinarily similar 2n = 14 karyotype in highly divergent marsupial lineages from both Australasia and the Americas provides strong evidence that it represents the ancestral marsupial karyotype. In other lineages, chromosome evolution has principally occurred by centric fissions and fusions as well as by changes in centromere position via inversions and centromeric transpositions. In multiple divergent marsupial families (e.g., Didelphidae, Macropodidae, Phalangeridae, Petauridae, Pseudocheiridae, Tarsipedidae) independent series of (mostly centric) fissions have led to increases in chromosome number from the ancestral 2n = 14 to 2n = 16–32. Subsequently in several lineages (e.g., Macropodidae, Potoroidae, Pseudocheiridae) these fissions have been followed by spectacular multiple series of (mostly centric) fusions, which in at least two cases has reduced the

Atlas of Mammalian Chromosomes, Edited by Stephen J. O’Brien, Joan C, Menninger, William G. Nash Copyright © 2006 John Wiley & Sons, Inc.

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MARSUPIALIA chromosome number to 2n = 10. Despite these often complex rearrangements, marsupial chromosomes have not been scrambled, and large blocks of the genome remain surprisingly intact. Evidence from G-banding and chromosome painting suggests that most marsupial karyotypes can be interrelated via the rearrangement of 15 conserved chromosomal segments. Compared to other mammals, marsupials have a relatively low number (2n = 10–32) of beautifully large chromosomes, although both the marsupial X and Y tend to be smaller than their eutherian (placental) counterparts. Because marsupial sex chromosomes lack a pseudoautosomal region, no homologous pairing, synaptonemal complex formation, or recombination occurs between the X and Y during male meiosis. However, comparative gene mapping and cross-species chromosome painting have shown that the marsupial X is homologous to large segments of the eutherian X chromosome, despite ~130 million years of independent evolution. Another unusual cytological feature of marsupials is the preferential inactivation of the paternally derived X chromosome in females and, in several species, the somatic elimination of sex chromosomes. Mark D. B. Eldridge C. J. Metcalfe

Order Didelphimorphia Four recent families of opossums are recognized within Didelphimorphia, all endemic to the Americas. Although this diverse order contains 16 genera and ~70 species, only three diploid numbers have been reported: 2n = 14, 18, 22.

Family Caluromyidae 2n = 14 (XX, XY) The family Caluromyidae (woolly opossums) contains two living genera and four species. All three members of the genus Caluromys have been karyotyped and two species have been G-banded. All possess a morphologically similar karyotype that can be related to the ancestral marsupial 2n = 14 complement by inversions in three or four autosomes. In two species (C. lanatus, C. philander) the (nucleolar organizing region) (NOR) is located on a single pair of autosomes.

Family Didelphidae 2n = 22 (XX, XY) This family of large opossums comprises 4 recent genera and 10 species, 9 of which have been karyotyped. All species are characterized by a similar 2n = 22 complement of graded acrocentric autosomes. However, there are some interspecies differences in X chromosome morphology as well as the presence and distribution of C-banding material. Representatives of three genera (Didelphis, Philander, Lutreolina) have been G-banded and all karyotypes show extensive homology. In addition, these G-banding patterns are consistent with the 2n = 22 didelphid karyotype being derived from the ancestral marsupial 2n = 14 complement via centric fissions in the four largest autosomes. The number and location of NORs varies within didelphids. In the yapok (Chironectes minimus) the NOR is located on one pair of autosomes, in both Philander and Lutreolina on two autosomal pairs, and in Didelphis on up to three pairs of autosomes.

ORDER DIDELPHIMORPHIA

Family Glironiidae Glironiidae is a monotypic family represented by the little known bushy-tailed opossum (Glironia venusta). No chromosome data are available.

Family Marmosidae 2n = 14, 18 (XX, XY) Although over 45% of species within this large family (~55 species) of mouse and short-tailed opossums have been karyotyped, only two chromosome numbers have been reported: 2n = 14 (Gracilinanus, Marmosa, Marmosops, Metachirus, Micoureus, Thylamys) and 2n = 18 (Monodelphis). All 2n = 14 species possess a karyotype morphologically similar to the ancestral marsupial complement, although the relative positions of some centromeres differ among species. There are also some interspecific differences in X chromosome size and morphology as well as the presence of C-banding material. However, G-banding data from representatives of five genera (Marmosa, Marmosops, Metachirus, Micoureus, Thylamys) have confirmed the basic homology of the marmosid 2n = 14 karyotype with the ancestral marsupial complement. These studies have shown that the marmosid 2n = 14 karyotype can be derived from the ancestral marsupial complement by as few as four changes in centromere position. Within 2n = 14 marmosids there is considerable variation in the number and location of NORs. In Gracilinanus, Metachirus, Thylamys, and Marmosops NORs have been reported from one pair of autosomes, in Micoureus from two autosomal pairs, and in Marmosa from up to three autosomal pairs. To date five species of Monodelphis have been karyotyped and all possess a similar 2n = 18 chromosomal complement. G-banding and crossspecies chromosome-painting data comparing the gray short-tailed opossum (M. domestica) with a range of other marsupial species have revealed extensive homology between chromosome arms. These data indicate that the 2n = 18 Monodelphis karyotype can be readily derived from the ancestral marsupial 2n = 14 complement by two fissions and several minor changes in centromere position. In Monodelphis, NORs have been reported from the short arms of chromosome 5 and the X. Mark D. B. Eldridge C. J. Metcalfe

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MARSUPIALIA

Marmosops incanus (South American Mouse Opossum)

2n=14

Svartman (1998)

Marmosa sp. (Mouse Opossum)

2n=14

Aniskin (1994a)

ORDER DIDELPHIMORPHIA

Micoureus demerarae (Mouse Opossum)

2n=14

Svartman (1998)

Thylamys elegans (Elegant Fat-Tailed Opossum/Southern Mouse Opossum)

2n=14

Shchipanov et al. (1996) Contributed by V. M. Aniskin

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14

MARSUPIALIA

Metachirus nudicaudatus (Brown Four-Eyed Opossum)

2n=14

Aniskin et al. (1991)

2n=14

Aniskin et al. (1991)

ORDER DIDELPHIMORPHIA

Monodelphis domestica (Short-Tailed Opossum)

2n=18

Graphodatsky, Perelman, Ferguson-Smith, Graves, and Ferquson (unpublished)

2n=18

Graphodatsky, Perelman, Ferguson-Smith, Graves, and Ferquson (unpublished)

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16

MARSUPIALIA

Ideogram: Monodelphis domestica (Short-Tailed Opossum)

2n=18

Pathak et al. (1993)

ORDER DIDELPHIMORPHIA

Caluromys lanatus (Woolly Opossum)

2n=14

Casartelli et al. (1986) Contributed by I. Ferrari

Philander opossum (Black Four-Eyed Opossum)

2n=22

Svartman (1998)

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18

MARSUPIALIA

Didelphis marsupialis (Common Opossum)

2n=22

Aniskin et al. (1991)

Didelphis virginiana (Virginia Opossum)

2n=22

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER PAUCITUBERCULATA

Order Paucituberculata This endemic South American order contains a single recent family.

Family Caenolestidae 2n = 14 (XX, XY) Seven living species of shrew opossum are divided into two genera, Caenolestes and Rhyncholestes. Although this family is poorly known, basic karyotypic data are available for five species, including representatives of both genera. All share a 2n = 14 karyotype that appears morphologically similar to the ancestral marsupial complement. However, there are some minor interspecific differences in autosomal centromere position and in X chromosome morphology. The Y chromosome is minute. Mark D. B. Eldridge C. J. Metcalfe

Rhyncholestes raphanurus (Chilean “Shrew” Opossum)

2n=14

Gallardo and Patterson (1987) Contributed by M. Gallardo

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20

MARSUPIALIA

Order Microbiotheria This South American order contains a single family and only one living species, the monito del monte (Dromiciops australis).

Family Microbiotheriidae 2n = 14 (XX, XY) The chromosomes of Dromiciops are morphologically similar to the 2n = 14 karyotype found in many other marsupials from both the Americas and Australasia. G-banding comparisons across marsupial orders have revealed extensive and remarkable homology and indicate that the Dromiciops karyotype can be derived from the ancestral marsupial 2n = 14 complement by three inversions. In Dromiciops, NORs are present on the two smallest autosomal pairs. The Y chromosome is minute. Although found only in southern South America, Dromiciops appears more closely related to Australasian marsupials rather to other American species. Mark D. B. Eldridge C. J. Metcalfe

Dromiciops australis (Monito del Monte)

2n=14(F)

2n=13(M)

Gallardo and Patterson (1987)

Gallardo and Patterson (1987)

Contributed by M. Gallardo

ORDER DASYUROMORPHIA

Order Dasyuromorphia Three recent families of carnivorous marsupials are recognized within Dasyuromorphia, all endemic to Australasia. Two families (Myrmecobiidae, Thylacinidae) are monotypic, while the third (Dasyuridae) is highly speciose.

Family Dasyuridae 2n = 14 (XX, XY) Dasyuridae is the largest marsupial family, containing 23 recent genera and ~68 species. Over 60% of known species have been karyotyped and representatives of at least 15 genera have been G-banded. All have been found to possess a virtually identical autosomal complement, with the NOR located terminally on the short arm of chromosome 5. Among species there is some minor variation in the amount and location of C-banding material as well as the morphology of the small X chromosome (~3% haploid genome). In all species the Y chromosome is punctiform. G-banding and reciprocal chromosome painting have revealed that the dasyurid 2n = 14 karyotype can be related to the ancestral marsupial 2n = 14 karyotype by six inversions.

Family Myrmecobiidae 2n = 14 (XX, XY) Myrmecobiidae includes only one known species, the numbat or banded anteater (Myrmecobius fasciatus). The numbat’s chromosomes appear similar in gross morphology to those of Dasyuridae. No further information is available.

Family Thylacinidae No data are available on the chromosomes of the now extinct thylacine or Tasmanian tiger (Thylacinus cynocephalus), which was the sole living representative of this family. However, it is tempting to speculate that, like all other members of Dasyuromorphia, it had a 2n = 14 complement. Mark D. B. Eldridge C. J. Metcalfe

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22

MARSUPIALIA

Antechinus flavipes (Yellow-Footed Marsupial “Mouse”/Antechinus)

2n=14

Rofe (1979)

2n=14

Rofe (1979) *Various banding levels

ORDER DASYUROMORPHIA

Planigale maculata (Flat-Skulled Marsupial “Mouse”/Planigale)

2n=14

Rofe (1979)

2n=14

Rofe (1979) *Various banding levels

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24

MARSUPIALIA

Ningaui sp. (Ningauis)

2n=14

Rofe (1979)

2n=14

Rofe (1979) *Various banding levels

ORDER DASYUROMORPHIA

Sminthopsis crassicaudata (Fat-Tailed Marsupial “Mouse”/Sminthopsis)

2n=14

Rofe (1979)

2n=14

Rofe (1979) *Various banding levels

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26

MARSUPIALIA

Antechinomys laniger (Kultarr)

2n=14

Rofe (1979)

2n=14

Rofe (1979) *Various banding levels

ORDER DASYUROMORPHIA

Dasyuroides byrnei (Kowari)

2n=14

Rofe (1979)

2n=14

Rofe (1979) *Various banding levels

27

28

MARSUPIALIA

Dasyurus hallucatus (Northern Quoll/Native “Cat”/Tiger “Cat”)

2n=14

Rofe (1979)

2n=14

Rofe (1979) *Various banding levels

ORDER DASYUROMORPHIA

Dasyurus viverrinus (Eastern Quoll/Native “Cat”/Tiger “Cat”)

2n=14

Rofe (1979)

2n=14

Rofe (1979) *Various banding levels

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MARSUPIALIA

Dasyurus maculatus (Spotted-Tailed Quoll)

2n=14

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Sarcophilus harrisii (Tasmanian Devil)

2n=14

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER PERAMELEMORPHIA

Order Peramelemorphia Peramelemorphia (bandicoots) contains two families and eight recent genera, all endemic to Australasia.

Family Peramelidae 2n = 14 (XX, XY); 2n = 19씹18씸 (XX, XY1Y2) Four genera and 10 recent species comprise the family Peramelidae. Three species are now extinct, but the remaining seven have been subject to cytogenetic analysis. With the exception of the greater bilby (Macrotis lagotis), all possess morphologically similar 2n = 14 karyotypes. G-banding of representatives from both Perameles (long-nosed bandicoots) and Isoodon (short-nosed bandicoots) show that these species retain the ancestral marsupial karyotype. In both Perameles and Isoodon the NOR is located on a single pair of autosomes. Members of these genera show sex chromosome mosaicism in many somatic tissues, with the inactivated paternally derived X chromosome being eliminated in females and the Y in males. In contrast the bilby does not show sex chromosome elimination and has a derived karyotype (2n = 19씹18씸) resulting from a series of fissions followed by several fusions.

Family Peroryctidae 2n = 14 (XX, XY) There are 3 genera and 11 species of rainforest bandicoots in the family Peroryctidae. This mainly New Guinean family is poorly known and only three species have been karyotyped. All possess a 2n = 14 karyotype morphologically similar to the ancestral marsupial complement, with the NOR on a single pair of autosomes. At least one species from both Echymipera and Microperoryctes shows sex chromosome mosaicism with one X chromosome being eliminated from some somatic tissues in females and the Y chromosome in males. In the common echymipera (E. kalabu) up to five supernumerary (or B) chromosomes can be present and these too are eliminated in some somatic tissues. Mark D. B. Eldridge C. J. Metcalfe

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32

MARSUPIALIA

Macrotis (Thylacomys) lagotis (Bilby/Rabbit-Eared Bandicoot)

2n=18 (F) 19 (M)

Martin and Hayman (1967) Contributed by D. Hayman

ORDER PERAMELEMORPHIA

Isoodon obesulus (Short-Nosed Bandicoot)

2n=14

Rofe (1979)

2n=14

Rofe (1979) *Various banding levels

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34

MARSUPIALIA

Perameles nasuta (Long-Nosed Bandicoot)

2n=14

Rofe (1979)

2n=14

Rofe (1979) *Various banding levels

ORDER NOTORYCTEMORPHIA

Order Notoryctemorphia This endemic Australian order contains two species, both placed in a single family and genus.

Family Notoryctidae 2n = 20 (XX, XY)? Found only in the deserts of central Australia, the fossorial marsupial moles are among the most elusive and enigmatic of mammals. Only a single individual male from the southern species (Notoryctes typhlops) has ever been karyotyped. In this specimen the submetacentric autosomes form a graded series, the fourth pair carrying a putative NOR. The large X chromosome (~6% haploid genome) is metacentric and the small Y acrocentric. In the absence of any additional studies, the relationship of the marsupial mole’s chromosomes to those of other marsupials remains unknown. Mark D. B. Eldridge C. J. Metcalfe

Notoryctes typhlops (Marsupial Mole)

2n=20

Calaby et al. (1974) Contributed by P. G. Johnston

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MARSUPIALIA

Order Diprotodontia Diprotodontia is the largest and most diverse marsupial order, containing 10 recent families. All ~140 diprotodont species are endemic to Australasia and include the koala, kangaroos, possums, and wombats. The entire spectrum of marsupial chromosome evolution is found within diprotodonts. Some species retain the 2n = 14 ancestral marsupial karyotype while others have highly derived karyotypes as a result of fissions, fusions, and other rearrangements. As a consequence, Diprotodontia contains species with both the highest (2n = 32) and lowest (2n = 10) known chromosome number for marsupials.

Family Acrobatidae 2n = 14 (XX, XY) The diminutive feather-tailed glider (Acrobates pygmaeus) and feather-tailed possum (Distoechurus pennatus) are the only living members of the Family Acrobatidae. Both species have 2n = 14 chromosomes similar to the ancestral marsupial complement, but differing slightly in the relative position of some centromeres. G-banding of Acrobates has revealed that its karyotype can be derived from the ancestral marsupial complement by 3 inversions.

Family Burramyidae 2n = 14 (XX, XY) Two genera of pygmy possums are recognized within Burramyidae: Burramys, which is monotypic, and Cercartetus, containing four species. All species share a similar 2n = 14 karyotype, although the mountain pygmy possum (Burramys parvus) differs in having three acrocentric chromosome pairs, in contrast to the distinctly biarmed complement of Cercartetus. G-banding data are only available for the western pygmy possum (C. concinnus), and its autosomes show a remarkable degree of homology to those of other 2n = 14 marsupial species, especially the common wombat (Vombatidae) and both the longnosed and short-nosed bandicoots (Peramelidae). These distantly related species are all believed to retain the ancestral marsupial karyotype. In Burramys and C. concinnus the NOR appears to be located on the X chromosome, while in the eastern (C. nanus) and little (C. lepidus) pygmy possums the NOR is located on different pairs of autosomes.

Family Macropodidae 2n = 10씸11씹 (XX, XY1Y2), 2n = 16씸15씹 (X1X1X2X2, X1X2Y), 2n = 12–24 (XX, XY) Macropodidae (kangaroos and wallabies) is one of the largest marsupial families containing 11 recent genera and 62 species. It is also the best studied cytologically with karyotypic data available for 83% of extant species. In all examined species the NOR is located on the X chromosome, although its position often varies between and sometimes within species. Representatives of most macropodid genera have been G-banded and many have also been examined with cross-species chromosome painting. Karyotypically this family is highly diverse, being characterized by two contrasting patterns—retention of an ancestral 2n = 22 karyotype in some lineages and multiple series of independent chromosome rearrangements in others. A 2n = 22 karyotype is considered ancestral for macropodids since a homologous complement is found in representatives of four genera (Petrogale, Thylogale, Setonix, Dorcopsis).

ORDER DIPROTODONTIA This 2n = 22 karyotype can be derived from the 2n = 14 ancestral marsupial complement by five fissions, one fusion, and several minor changes in centromere position. All other macropodid species can be readily derived from this 2n = 22 karyotype principally via centric fusions. For example, the red kangaroo (Macropus rufus) has a 1–10 fusion (2n = 20); the grey kangaroos (M. giganteus, M. fuliginosus) 1–8, 5–9, and 6–10 fusions (2n = 16); the wallaroo (M. robustus) 1–10, 5–6, and 8–9 fusions (2n = 16); and the tammar wallaby (M. eugenii) 1–10, 5–8, and 6–9 fusions (2n = 16). In addition, similar independently derived fusions are known from six other macropodid genera. In two species the fusion events have involved the sex chromosomes. The swamp wallaby (Wallabia bicolor) is characterized by fusions between ancestral macropodid chromosomes 1–9, 4–6–10, 5–8, and 2–7–X, resulting in a 2n = 10 complement in females and 2n = 11 in males. In the spectacled hare wallaby (Lagorchestes conspicillatus) fusions have occurred between ancestral chromosomes 6–10–7 as well as between chromosome 3 and both X and Y. Subsequently the 3–Y has also fused to chromosome 8, giving a 2n = 15 complement in males but 2n = 16 in females. Among macropodid species there is considerable variation in the amount and location of C-banding material, although, when present, it is usually centromeric. In addition, the size (~2–12% haploid genome) and morphology of the X chromosome are often variable among and sometimes within species.

Family Petauridae 2n = 18, 20, 22 (XX, XY) Chromosome data are available for 6 of the approximately 10 known petaurid species. The sugar (Petaurus breviceps), squirrel (P. norfolcensis), and yellow-bellied (P. australis) gliders all possess a similar 2n = 22 karyotype of metacentric and submetacentric chromosomes. A G-banding comparison of P. norfolcensis with other marsupials has identified limited regions of homology, suggesting that the derivation of the Petaurus karyotype has been, for marsupials, unusually complex. The relationship of the Petaurus complement to the morphologically distinct karyotypes found in Leadbeater’s possum (Gymnobelideus leadbeateri), 2n = 22, and the striped possums (Dactylopsila spp.), 2n = 18, is unkown.

Family Phalangeridae 2n = 14, 20 (XX, XY) The chromosomes of cuscuses and brushtail possums are poorly studied, with diploid numbers reported for only 7 of the 21 known species. In addition, Gbanding and cross-species chromosome-painting data are available for only a single species, the common brushtail (Trichosurus vulpecula), 2n = 20. These data indicate that the Trichosurus karyotype can be readily derived from the 2n = 14 ancestral marsupial karyotype by four centric fissions, one fusion, and several minor changes in centromere position. However, the relationship of the Trichosurus karyotype with the morphologically distinct 2n = 14 complement found in cuscuses (Phalanger spp.) remains unexplored.

Family Phascolarctidae 2n = 16 (XX, XY) An Australian faunal icon, the koala (Phascolarctos cinereus) is the only modern representative of the family Phascolarctidae. G-banding studies have shown that the koala’s karyotype can be simply derived from the 2n = 14 ances-

37

38

MARSUPIALIA tral marsupial complement by a single centric fission and one minor change in centromere position.

Family Potoroidae 2n = 12씸13씹 (XX, XY1Y2), 2n = 22, 24, 34 (XX, XY) Potoroidae contains 11 recent species of rat-kangaroo placed into 5 genera. Two species are now extinct but chromosome data are available for the remaining 9. In all species the NOR is located on the X chromosome. The musky rat-kangaroo (Hypsiprymnodon moschatus) and four Bettongia species have a 2n = 22 karyotype, although in Bettongia the chromosomes are more metacentric. The relationship between the potoroid 2n = 22 karyotype and the ancestral marsupial (2n = 14) or the ancestral macropodid (2n = 22) karyotype is currently unknown. G-banding and/or cross-species chromosome-painting data are available for all three species of potoroo (Potorous spp.). The 2n = 24 karyotype of the long-footed potoroo (P. longipes), which may be ancestral for potoroids, shows extensive homology to the ancestral macropodid 2n = 22 karyotype, from which it can be derived by a single centric fission and one change in centromere position. The karyotypes of the other two species (P. tridactylus, P. gilberti), both identical 2n = 12씸13씹, can be readily derived from the P. longipes karyotype by six fusions (mostly centric and including an autosome to X translocation) and one change in centromere position. The opposite trend (i.e., fissions) is apparent in the karyotype of the rufous bettong (Aepyprymnus rufescens), 2n = 32. Cross-species chromosome painting suggests that the Aepyprymnus karyotype can be derived from the 2n = 24 complement of P. longipes by four noncentric fissions.

Family Pseudocheiridae 2n = 10–22 (XX, XY) Chromosome data for this cytogenetically interesting family are unfortunately limited. Of the 16 known ringtail possum species, diploid numbers have been reported for 10, and G-banding data are available for only 2 species. In addition, the relationship between pseudocheirid chromosomes and those of other marsupials remains unknown. Two Australian species, the greater glider (Petauroides volans) and the lemouroid ringtail (Hemibelideus lemuroides), have been well studied cytogenetically. Both species show sex chromosome mosaicism, with the Y being eliminated from most somatic tissues in males of both species and one X also being eliminated in H. lemuroides females. The greater glider is also of interest since individuals can possess up to eight supernumerary (or B) chromosomes. G-banding data for the greater glider (2n = 22) and the lemouroid ringtail (2n = 20) indicate that they differ by a single fission/fussion event. The NOR is located on the X in P. volans and autosomally in H. lemuroides. In the common ringtail (Pseudocheirus peregrinus), 2n = 20, NORs may be present on up to three pairs of autosomes and the sex chromosomes. This species also shows considerable diversity in chromosome size and morphology due to the variable presence of centromeric constitutive heterochromatin. In the Herbert River ringtail (Pseudochirulus herbertensis), 2n = 12, and the closely related Daintree River ringtail (Pseudochirulus cinereus), 2n = 16, NORs are found on two pairs of autosomes and these species differ by two centric fusion/fission events. In addition, both species share enormous sex chromosomes (X ~16%, Y ~12% haploid genome), which appear to result from the independent fusion of the X and Y to a homologous pair of autosomes.

ORDER DIPROTODONTIA

Family Tarsipedidae 2n = 24 (XX, XY) The honey possum (Tarsipes rostratus) is the only known member of the family Tarsipedidae. Tarsipes has an all-acrocentric karyotype with NORs present on the X and two pairs of autosomes. G-banding studies have shown that the honey possum’s chromosomes can be readily derived the 2n = 14 ancestral marsupial complement by five fissions (four centric) and several minor changes in centromere position.

Family Vombatidae 2n = 14 (XX, XY) The three recent species of wombats are placed in two genera (Vombatus and Lasiorhinus), although all possess morphologically similar karyotypes. Gbanding comparisons suggest that the common wombat (V. ursinus) retains the ancestral marsupial karyotype, from which the southern hairy-nosed wombat (L. latifrons) differs by inversions in the first two pairs of autosomes. Cross-species chromosome painting has shown that the chromosomes of L. latifrons differ from the 2n = 14 ancestral marsupial complement by two inversions in chromosome 1. In L. latifrons, NORs are located on the X chromosome and two pairs of autosomes, while in Vombatus a putative NOR has been identified on the X chromosome. Mark D. B. Eldridge C. J. Metcalfe

39

40

MARSUPIALIA

Phascolarctos cinereus (Koala)

2n=16

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER DIPROTODONTIA

Vombatus ursinus (Common Wombat/Coarse-Haired Wombat)

2n=14

Rofe (1979)

2n=14

Rofe (1979) *Various banding levels

41

42

MARSUPIALIA

Lasiorhinus latifrons (Northern Hairy-Nosed Wombat/ Soft-Furred Wombat)

2n=14

Rofe (1979)

2n=14

Rofe (1979) *Various banding levels

ORDER DIPROTODONTIA

Trichosurus vulpecula (Common Brush-Tailed Possum)

2n=20

Rofe (1979)

2n=20

Rofe (1979) *Various banding levels

43

44

MARSUPIALIA

Potorus tridactylus (Long-Nosed Potoroo)

2n=12

Graphodatsky, Perelman, Rens, Ferguson-Smith, Graves, and Ferguson (unpublished) Contributed by A. Graphodatsky

ORDER DIPROTODONTIA

Thylogale billardierii (Red-Bellied Pademelon)

2n=22

Rofe (1979)

2n=22

Rofe (1979) *Various banding levels

45

46

MARSUPIALIA

Thylogale thetis (Red-Necked Pademelon)

2n=22

Eldridge et al. (1992a)

Petrogale xanthopus celeris (Yellow-Footed Rock Wallaby)

2n=22

Eldridge et al. (1992a)

ORDER DIPROTODONTIA

Petrogale xanthopus xanthopus (Yellow-Footed Rock Wallaby)

2n=22

Eldridge et al. (1992a)

Petrogale lateralis lateralis (Black-Footed Rock Wallaby)

2n=22

Eldridge et al. (1992a)

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48

MARSUPIALIA

Petrogale penicillata (Brush-Tailed Rock Wallaby)

2n=22

Rofe (1979)

2n=22

Rofe (1979) *Various banding levels

ORDER DIPROTODONTIA

Petrogale concinna (Little Rock Wallaby)

2n=16

Eldridge et al. (1992b)

Petrogale brachyotis (Short-Eared Rock Wallaby)

2n=20

Eldridge et al. (1992b)

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50

MARSUPIALIA

Wallabia bicolor (Swamp Wallaby)

2n=10(F) 11(M)

Rofe (1979)

2n=10(F) 11(M)

Rofe (1979)

ORDER DIPROTODONTIA

Macropus parma (Parma Wallaby)

2n=16

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Macropus rufogriseus (Red-Necked Wallaby)

2n=16

Rofe (1979)

51

52

MARSUPIALIA

Macropus eugenii (Tammar Wallaby)

2n=16

Graphodatsky, Perelman, Rens, Ferguson-Smith, Graves, and Ferguson (unpublished) Contributed by A. Graphodatsky

2n=16

Graphodatsky, Perelman, Rens, Ferguson-Smith, Graves, and Ferguson (unpublished) Contributed by A. Graphodatsky

ORDER DIPROTODONTIA

Macropus eugenii (Tammar Wallaby)

2n=16

Rofe (1979) *Various banding levels

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54

MARSUPIALIA

Macropus parryi (Whiptail Wallaby)

2n=16

Rofe (1979)

2n=16

Rofe (1979) *Various banding levels

ORDER DIPROTODONTIA

Macropus giganteus (Eastern Gray Kangaroo)

2n=16

Rofe (1979)

2n=16

Rofe (1979) *Various banding levels

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56

MARSUPIALIA

Macropus fuliginosus (Western Gray Kangaroo)

2n=16

Rofe (1979)

2n=16

Rofe (1979) *Various banding levels

ORDER DIPROTODONTIA

Macropus robustus (Common Wallaroo)

2n=16

Rofe (1979)

2n=20

Rofe (1979)

2n=20

Rofe (1979) *Various banding levels

Macropus rufus (Red Kangaroo)

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58

MARSUPIALIA

Cercartetus concinnus (Western Pygmy Possum)

2n=14

Rofe (1979)

2n=14

Rofe (1979) *Various banding levels

ORDER DIPROTODONTIA

Pseudocheirus peregrinus (Common Ring-Tailed Possum) P. p. cookii

P. p. puicher

P. p. rubidus

2n=20

Murray et al. (1980)

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60

MARSUPIALIA

Petropseudes dahli (Rock Ring-Tailed Possum)

2n=16

Murray et al. (1980)

2n=16

Murray et al. (1980)

Hemibelideus lemuroides (Lemuroid Ring-Tailed Possum/Greater Gliding Possum)

2n=20

McQuade (1984)

ORDER DIPROTODONTIA

Petaurus norfolcensis (Lesser Gliding Possum/Squirrel Glider)

2n=21

Rofe (1979)

Tarsipes rostratus (Honey Possum)

2n=24

Hayman (1990)

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62

MARSUPIALIA

Acrobates pygmaeus (Pygmy Gliding Possum/Feather-Tailed Glider)

2n=14

Rofe (1979)

2n=14

Rofe (1979) *Various banding levels

AFROTHERIA Order Afrosoricida Order Macroscelidea Introduction to Golden Moles, Elephant Shrews, and Tenrecs The grouping of golden moles, tenrecs, and elephant shrews (termed Afroinsectivora or African insectivores by Waddell et al., 2001) within Afrotheria* is one of the more dramatic departures from the morphologically based evolutionary tree to have emerged from recent molecular phylogenetic investigations (e.g., Murphy et al., 2001a,b; Amrine-Madsen et al., 2003; Novacek, 1986 and references therein). In the past, golden moles and tenrecs were considered part of Lipotyphla (formerly Insectivora), a purportedly monophyletic group comprising shrews (Soricidae), hedgehogs (Erinaceidae), moles (Talpidae), and solenodons (Butler, 1988) and their exclusion from Lipotyphla is in contradiction to more than a century of morphological inference (e.g., Haeckel, 1866; Gregory, 1910; Butler, 1988; MacPhee and Novacek, 1993). Likewise, elephant shrews have conventionally been grouped together with the glirids (lagomorphs and rodents) in Anagalidia, and their suggested affiliation with a tenrec-golden mole clade Afrosoricida (M.S. Stanhope et al., 1998) within Afroinsectiphillia (aardark + Afroinsectivora) is novel. The species taxonomy adopted below for Afroinsectivora follows that of the IUCN’s (International Union for Communication of Nature) Afrotherian Specialist Group (http:// www.calacademy.org/research/bmammals/afrotheria/systematics.html). Cytogenetic information on the three afroinsectivoran groups is relatively sparse. Of the 21 species of golden moles (family Chrysochloridae), conventionally banded karyotypes are available for 12 species representing 6 of 9 recognized genera (Amblysomus, Neamblysomus, Calcochloris, Chlorotalpa, Chrysochloris, and Chrysospalax). G-bands are available only for the Cape golden mole, Chrysochloris asiatica (2n = 30; Robinson et al., 2004). Known chromosome numbers range from 2n = 28 (Calcochloris obtusirostris) to 2n = 36 in Amblysomus robustus (Bronner, 1995b). It is noteworthy that Bronner (1995a) originally regarded A. hottentotis as comprising three allopatric cytotypes (2n = 30, 34, 36) but the 2n = 34 cytotype was subsequently described as a valid species, A. septentrionalis (Bronner, 1996), with the 2n = 36 form being assigned to A. robustus (Bronner, 1995b). All other species have 2n = 30. *A supraordinal grouping of African mammals including elephants, hyraxes, dugongs manatees, golden moles, elephant shrews, tenrecs, and aardvark (Springer et al., 1997; Stanhope et al., 1998; Robinson and Seifert, in press, and references therein). Atlas of Mammalian Chromosomes, Edited by Stephen J. O’Brien, Joan C. Menninger, William G. Nash Copyright © 2006 John Wiley & Sons, Inc.

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64

AFROTHERIA Variation in diploid number among the elephant shrews (family Macroscelidea) is equally constrained. Of the 15 recognized species, seven have been karyotyped representing all but one (Rhynchocyon) of the recognized genera. G-banded chromosomes have been reported for Elephantulus rupestris (2n = 26; Wenhold and Robinson, 1987; Robinson et al., in press), Macroscelides proboscideus (2n = 26; Wenhold and Robinson, 1987; Svartman et al., 2004), and Petrodromus tetradactylus (2n = 28; Wenhold and Robinson, 1987). Data from unbanded preparations exist for Elephantulus edwardii (2n = 26; Tolliver et al., 1989), E. myurus (2n = 30; Ford and Hamerton, 1956; Tolliver et al., 1989), E. brachyrhynchus (2n = 26; Stimson and Goodman, 1966; Tolliver et al., 1989), and E. intufi (2n = 26; Tolliver et al., 1989). The kinds of rearrangements underpinning what little chromosomal variation there is among the elephant shrews is not clear, although Wenhold and Robinson (1987) documented a tandem fusion difference between the 2n = 28 of P. tetradactylus and the 2n = 26 of M. proboscideus and E. rupestris. Tenrecs (family Tenrecidae, 30 species), unlike the golden moles and elephant shrews, have only marginal representation on continental Africa, being restricted almost exclusively to Madagascar. Of the 10 recognized genera only otter shrews (Potomogalinae; Micropotamogale and Potamogale) are found on the African continent. Known diploid numbers vary from 2n = 14 (Geogale aurita; Olsen, personal communication) to 2n = 40 (Echinops telfairi; Benirschke, 1969). Unbanded karyotypes (Bernischke, 1969) are available for E. telfairi, Hemicentetes nigriceps (2n = 38), and Micropotamogale dobsoni (2n = 30). Morphological and molecular studies have suggested conflicting phylogenetic relationships within elephant shrews (Corbet and Hill, 1992 and Corbet, 1954; Douady et al., 2003) and tenrecs (Asher, 2001; Douady et al., 2002), in part reflecting poor taxon sampling. The only phylogenetic information on golden moles is that of Bronner (1995a) based on craniodental characters among 10 chyrsochlorid species from South Africa. Clearly, the presence of conserved karyotypes in most Afroinsectivora species limits their usefulness in refining relationships within the various assemblages. However, recent cross-species chromosome-painting studies on elephant shrews and golden moles have provided further support for the monophyly of Afrotheria (Svartman et al., 2004; Robinson et al., 2004) and, together with data on the aardvark (Yang et al., 2003a), on some of the problematic associations within Afroinsectivora. Using human (HSA) chromosome-specific painting probes, Robinson et al. (2004) identified one segmental configuration (HSA 2/8p/4) in the golden mole, elephant shrew, and aardvark that is absent in the elephant (Frönicke et al., 2003; Yang et al., 2003a), thus supporting the recognition of Afroinsectiphillia. Additionally, although more weak, their painting data suggest that elephant shrews and aardvark are sister taxa (the syntenic associations HSA 10q/17 and HSA 3/20 are present in both the elephant shrew and aardvark but absent in the golden mole and elephant). They failed to find evidence in support for an elephant shrew–golden mole affiliation within Afroinsectivora (tenrecs were not included in their study). In conclusion Afroinsectivora species, with the exception of tenrecs, are characterized by extreme karyotypic conservation. Their low diploid numbers, basal position in the eutherian evolutionary tree, and large number of primitive syntenic associations identified by cross-species chromosome painting (Svartman et al., 2004; Robinson et al., 2004) all suggest that they have retained a karyotype that largely resembles that of the last common ancestor of Placentalia. Terrence J. Robinson

ORDER AFROSORICIDA

Chlorotalpa duthieae (African Golden Mole)

2n=30

Bronner (1995a)

Chlorotalpa sclateri (African Golden Mole)

2n=30

Bronner (1995a)

65

66

AFROTHERIA

Amblysomus julianae (South African Golden Mole)

2n=31

Bronner (1995a)

Amblysomus iris (South African Golden Mole)

2n=34

Capanna et al. (1989) Contributed by E. Nevo

ORDER AFROSORICIDA

Amblysomus hottentotus (South African Golden Mole)

2n=30

Bronner (1995a)

Amblysomus hottentotus (South African Golden Mole)

2n=36

Bronner (1995a)

67

68

AFROTHERIA

Microgale dobsoni (Long-Tailed Tenrec/Shrewlike Tenrec)

2n=30

Benirschke (1969)

Hemicentetes nigriceps (Streaked Tenrec)

2n=38

Benirschke (1969)

ORDER AFROSORICIDA

Echinops telfairi (Small Madagascar Hedgehog)

2n=40

Benirschke (1969)

69

70

AFROTHERIA

Order Macroscelidea

Macroscelides proboscideus (Short-Eared Elephant Shrew)

2n=26

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER SIRENIA

Order Sirenia The sirenians are a group of two recent families, Dugongidae (dugongs and sea cows) and Trichechidae (manatees). Along with elephants and hyraxes, they comprise the Paeungulata superorder of herbibore species within the Afrotheria mammalian superorder. There are four living species of Sirenia today: dugong, West Indian manatee, West African manatee, and Amazonian manatee. A fifth modern dugong species, the Steller’s sea cow, formerly ranged from Baja California north along the Pacific coast of North America to the Aleutian Islands as recently as 100,000 years ago. Unable to submerge, the sea cows used their appendages to support themselves against rocks to feed on kelp. An easy target for human exploitation, this species was driven to extinction by 1768. Sirenians are large sea mammals that feed on water plants in shallow tropical coastal waters, bays, estuaries, lagoons, and rivers in the Old and New World. They evolved from terrestrial mammals akin to elephants and hyraxes that once browsed the grassy coastal swamps of the Paleocene some 60 mya. Sirenians have a rounded torpedo-shaped body, smallish heads, fore-limb flippers, and tail fins adapted for sea life. The order Sirenia numbers fewer than 60,000 individuals surviving today, the lowest of any mammalian order. Manatees are listed as vulnerable by the IUCN, endangered by the USFWS (U.S. Fish & Wildlife Service), and Appendix I by CITES (Convention on International Trade in Endangered Species). West Indian manatess were estimated at around 1200 off the Florida coast, but most populations are fewer than 100 animals. Nearly all recovered animals show scars from motorboat propellers. We present here the karyotype of the Florida manatee, Trichechus manatus (Gray et al., 2002). Stephen J. O’Brien

71

72

AFROTHERIA

Trichechus manatus latirostris (Florida Manatee)

2n=48

Gray et al. (2002)

ORDER SIRENIA

Ideogram: Trichechus manatus latirostris (Florida Manatee)

2n=48

Gray et al. (2002)

Sex chromosomes

73

74

AFROTHERIA

Order Proboscidea Elephant Cytogenetics Elephants are the largest living terrestrial animals. Three living species comprise the order Proboscidea, which also includes many extinct forms. The two living genera, which diverged 5–7 mya, comprise the family Elephantidae (Maglio, 1973; Vignaud et al., 2002). The Asian elephant, Elephas maximus, is the only living species in its genus. It is endangered with a surviving population of 35,000–50,000 across 14 countries in South and Southeast Asia (Kemf and Santiapillai, 2000). The African genus Loxodonta had also been considered monotypic, but recent morphological and genetic studies have demonstrated that forest and savanna African elephants are distinct species, Loxodonta africana (African savanna elephant) and Loxodonta cyclotis (African forest elephant). Morphometric analyses of 295 elephant skulls from across the continent found that African elephants form two morphologically distinct species with few intermediate phenotypes (Groves and Grubb, 2000; Grubb et al., 2000). Genetic analyses using both nuclear gene sequences and microsatellites found deep genetic separation between forest and savanna elephants, estimated at 2.6 million years (Roca et al., 2001; Comstock et al., 2002). The two African species hybridize where their habitats meet, but the genetic integrity of the parent species appears intact, likely due to selection against hybrid phenotypes (Roca et al., 2001). The forest elephant, L. cyclotis, inhabits tropical forests in West and Central Africa, while the savanna or bush elephant, L. africana, is found in a variety of habitats surrounding the tropical forests. Mitochondrial deoxyribonucleic acid (DNA) sequences had suggested that West African elephants may be genetically distinctive (Eggert et al., 2002). This finding has been questioned (Debruyne et al., 2003; Roca et al., 2005) and awaits verification using nuclear genes, while morphological studies of West African elephants have proven inconclusive (Groves, 2000). Across Africa, a total of 600,000 elephants survive in 37 African nations (Blanc et al., 2003).

Family Elephantidae 2n = 56 The diploid chromosome number is 56 for both African savanna and Asian elephants, as espoused by all published reports (Hungerford et al., 1966; Norberg, 1969; Hosli and Thurig, 1970; Thurig, 1970; Wallace, 1988; Groves, 2000; Houck et al., 2001; Fronicke et al., 2003; Yang et al., 2003a). Elephant karyotypes have been analyzed by chromosome G-banding only recently by Houck et al. (2001), with similar results reported by Fronicke et al. (2003). A high level of chromosome band homology was found between African savanna and Asian elephants (Houck et al., 2001). Of 27 autosome pairs, 26 were conserved between the genera (Hungerford et al., 1966; Houck et al., 2001). Loxodonta africana had 25 acrocentric/telocentric and 2 metacentric/ submetacentric autosomal pairs, while Elephas maximus had one less acrocentric and one more submetacentric due to either heterochromatic arm addition or deletion involving autosomal pair 27 (Houck et al., 2001). Several acrocentric autosomes in L. africana had small short arms that were absent in E. maximus (Houck et al., 2001). The X chromosome is large and submetacentric (Hungerford et al., 1966; Houck et al., 2001). The Y chromosome is small and acrocentric in L. africana and slightly larger in E. maximus with more distinct G-bands (Houck et al., 2001). Two reports of human

ORDER PROBOSCIDEA chromosome-painting probes on elephants agreed overall except for the association of human chromosomes HSA 5 and 21 to elephant chromosome LAF 3 (Fronicke et al., 2003; Yang et al., 2003a). The flow karyotype was different across the two studies, likely due to polymorphisms of the heterochromatic short arms of the acrocentric elephant chromosomes (Fronicke et al., 2003). Interordinal chromosome painting using human paint probes on African savanna elephant chromosomes identified 53 evolutionary conserved segments on 28 chromosomes (27 autosomes and X chromosome) (Fronicke et al., 2003). Reciprocal experiments using probes derived from flow-sorted African savanna elephant chromosomes identified 68 conserved segments in the human nuclear genome (Fronicke et al., 2003). Chromosome-painting studies on African savanna elephants have shed light on the early evolution of eutherian chromosomes. The closest relatives to Proboscidea are the orders Sirenia (manatees and dugongs) and Hyracoidea (hyraxes); the three along with extinct orders are collectively known as Paenungulata and form a group within Afrotheria, a superordinal clade of eutherian mammals that also includes aardvarks, elephant shrews, tenrecs, and golden moles (Springer et al., 1997). Afrotherians diverged from other eutherians 107 mya following tectonic isolation of the Afro-Arabian continent (Springer et al., 1997, 2003). The first nonsequencing support for the clade Afrotheria was provided by in-situ hybridization of human chromosome paints, which detected chromosomal arrangements unique to the first two afrotheres examined: elephant and aardvark (Fronicke et al., 2003; Yang et al., 2003a). The afrotherian arrangements are the association of human chromosomes HSA 5 and 21 to elephant chromosome LAF 3 and HSA 1 and 19 to LAF 2 (Fronicke et al., 2003). These associations were later confirmed for a third order within Afrotheria, Macroscelidea, the elephant shrews (Svartman et al., 2004). Recent genetic analyses may be used to identify additional elephant populations as candidates for intraspecific cytological comparisons. Only mainland Asian elephants have been used for cytogenetic studies (Hungerford et al., 1966), and a lack of intraspecific differences had been noted between an elephant from India and another from Thailand (Hungerford et al., 1966). However the islands of Sumatra and Borneo have genetically distinctive populations (Fleischer et al., 2001; Fernando et al., 2003), suggesting the two island populations as potential candidates for intraspecific cytologenetic studies. African savanna elephants show little population genetic divergence continentwide, although Cameroon elephants are somewhat different from the much larger populations of eastern and southern savanna elephants (Comstock et al., 2002). However, all savanna elephants used for cytological studies have been from eastern (Hungerford et al., 1966; Thurig 1970; Fronicke et al., 2003) or southern (Wallace, 1988) Africa. The African forest elephant has never been karyotyped (Fronicke et al., 2003), due to its absence from Western zoos. The forest species is thus an interesting prospect for cytogeneticists. Forest elephants display remarkable intraspecific nucleotide diversity that is 29 times higher than detected among savanna elephants (Roca et al., 2001). The very high levels of polymorphism in forest elephants may be due to the isolation of populations in allopatry during the Pleistocene ice ages, when the forests of Africa diminished in range to form isolated refugia, followed by mixing of the populations (Wahlund effect) as forests became contiguous following the ice ages (Behrensmeyer, 1992; Roca et al., 2001). Although genetic polymorphism is not indicative of chromosome rearrangements and African savanna and Asian elephant karyotypes are quite similar (Houck et al., 2001), it would nonetheless be worthwhile to examine forest

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AFROTHERIA elephants for potential cytogenetic polymorphisms. Since forest elephants in the Guinean and Congolian forest zones are separated by the unforested Benin or Dahomey gap (White, 1983), examination of forest elephants on both sides of the gap may prove valuable. Alfred L. Roca

Elephas maximus (Asian/Indian Elephant)

2n=56

Houck et al. (2001)

ORDER PROBOSCIDEA

Loxodonta africana (African Savanna Elephant)

2n=56

Houck et al. (2001)

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Order Hyracidea The order Hyracoidea comprises a single family of herbivorous rabbit-sized mammals, Procaviidae. The range of hyraxes is confined to rocky terrains and savanna habitats of Africa and southwest Asia. Classical morphological studies identified Proboscidea (elephants) and Sirenia (manatees and see cows) as the closest living relatives of Hyracoidea, an observation which is confirmed by recent comprehensive DNA sequence comparisons (Murphy et al., 2001a,b; Madsen et al., 2001). The up to 14 hyrax species are grouped into three genera: Procavia (rock hyraxes or dassies), Heterohyrax (yellow-spotted hyraxes), and Dendrohyrax (tree or bush hyraxes). The karyotypic data for Hyracoidea are limited to three species each representing one genus: Procavia capensis (unbanded karyotypes: Hungerford and Snyder, 1969, Hsu and Benirschke, 1971; GTG-banding: the karyotype presented here), Dendrohyrax arboreus (GTG-banding and C-banding: Prinsloo and Robinson, 1991), and Heterohyrax brucei (GTG-banding and C-banding: Prinsloo and Robinson, 1991). Each of the karyotypes displays a chromosome number of 2n = 54, which was also suggested to be the ancestral Hyracoidea chromosome number. The karyotypes consist predominantly of acrocentric chromosome pairs (H. brucei: 20, P. capensis: 19, D. arboreus: 15). Prinsloo and Robinson (1991) were able to identify G-banding homologies for 21 out of 26 autosomes between D. arboreus and H. brucei. However, the two karyotypes showed other remarkable differences, which could be mainly explained by the addition of heterochromatic short arms and terminal C-bands in D. arboreus. Unfortunately, the print reproduction of these karyotypes is very poor, so that comparisons to the G-banded chromosomes of the rock hyrax (see below) are very difficult. Thus, unambiguous homologies can only be identified for P. capensis chromosomes 1, 2, and 4 to chromosomes 1, 2, and 4 of D. arboreus, respectively. The X chromosome morphology has been conserved between the three hyrax species. Whereas the opportunities for comparative analyses inside Hyracoidea are limited, comparisons to elephant karyotypes (Houck et al., 2001) reveal great similarities in banding patterns between P. capensis chromosomes 4, 7, 8, and 10 and African elephant (Loxodonta africana) chromosomes 6, 8, 17, and 10, respectively. Additionally, rock hyrax chromosomes 2 and 3 might be derived from elephant chromosomes 2 and 1 by inversions. Thus the karyotype studies provide another indication for the phylogenetic relationship of such diverse clades as elephant and hyraxes. The hyrax specimen was kindly made available by Dr. Bert Geyer (Zoo Frankfurt, Frankfurt, Germany). Lütz Froenicke

ORDER HYRACIDEA

Procavia capensis (Hyrax/Rock Dassie)

2n=54

Fronicke (unpublished)

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Order Tubulidentata The order Tubulidentata contains one family, Orycteropodidae, with a single living genus and species, Orycteropus afer, which is distributed in Africa south of the Sahara (Nowak, 1999). Aardvarks were once grouped with the anteaters, sloths, and armadillos and pangolins in the order Edentata. Recent molecular phylogenetic studies place the aardvark firmly in the superordinal clade Afrotheria, which includes the hyraxes, sirenians, elephants, tenrecs, golden moles, and elephant shrews (Springer et al., 1997; M. J. Stanhope et al., 1998b; Murphy et al., 2001a,b; Madsen et al., 2001). Descriptions are available on the conventional (Benirschke et al., 1970) and G- and C-banded karyotypes (Pathak et al., 1980). The aardvark has a 2n = 20 karyotype that consists of two pairs of large subtelocentric chromosomes plus seven pairs of medium to small metacentric autosomes and the sex pair. Most recently, a genomewide comparative chromosome map between aardvark and human has been established by cross-species reciprocal painting, demonstrating that the aardvark has a highly conserved karyotype which has retained almost all the ancestral conserved syntenies identified in eutherian mammals (Yang et al., 2003a). Fentang Yang

Orycteropus afer (Aardvark/Ant Bear)

2n=20

Yang et al. (2003a)

XENARTHRA Order Xenarthra A fascinating exhibit in the Ameghino Museum of Natural History of La Plata, Argentina, shows the large variety of poorly understood members of this order beneath the shell of a glyptodont, one of the ancestors. One can readily recognize the shell markings of modern armadillos on this large overarching shell. Indeed, these are fascinating animals and a wonderful symposium describing their biology was edited by Montgomery (1985). Xenarthra (formerly Edentata) contains odd-jointed mammals that date from about 100 mya and they are basically South American animals. Xenarthra is comprised of two major groups, those species with shells and those with hair, and we recognize 29 species in four families (but different schemes of taxonomy have also been debated): Bradypodidae, or three-toed sloths, possibly contains as many as 3 species; Megalonychidae, or two-toed sloths, 2 species; Myrmecophagidae, or anteaters, 4 species; and Dasypodidae, or armadillos, 20 species. The variety of the latter group is rarely appreciated since the nine-banded armadillo, a recent immigrant to North America, is often thought to be the most representative of this group. In contrast to several molecular genetic studies (e.g., Delsuc et al., 2002), regrettably few karyotypic studies have been conducted in these animals and none have been carried out with the most modern analytical modalities. In part this may be the result of their often nocturnal habits and thus their usual absence from most zoological gardens. They are thus not readily accessible for cytogenetic study, the nine-banded armadillo being the sole exception. This species and the “mulita” also have the unusual feature of polyembryony (multiple identical offspring) and a single uterus.

Bradypodidae Jorge and his collaborators (Jorge et al., 1977, 1985) are the only investigators to have studied two of these species. They found between 52 and 55 chromosomes from animals at different locations, nearly all elements being acrocentrics, except for a metacentric X.

Megalonychidae Choloepus has presented problems in karyotyping. Both species have been studied but with very variable results. Animals from Manaus had 64 chromoAtlas of Mammalian Chromosomes, Edited by Stephen J. O’Brien, Joan C. Menninger, William G. Nash Copyright © 2006 John Wiley & Sons, Inc.

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XENARTHRA somes; others from the same and other South American locations had between 49 and 53 elements. Most chromosomes are acrocentric. The structure and nature of the sex chromosomes, however, are very poorly understood. Gbanding and some C-banding studies exist.

Myrmecophagidae Tamanduas or lesser anteaters have 54 chromosomes, mostly metacentric, while the giant anteater carries 60 chromosomes with many more acrocentric.

Dasypodidae Nine-banded (Beath et al., 1962) and seven-banded armadillos (Dasypus) possess 64 mostly acrocentric chromosomes. Six-banded (Euphractus) armadillos have 58 chromosomes, hairy armadillos have 2n = 60, and softtailed (Cabassous) possess 62 elements. The giant armadillo (Benirschke et al., 1969; Benirschke and Wurster, 1969) (Priodontes), perhaps the rarest and most primitive-appearing species, has a complement of 50 chromosomes, about half of which are acrocentric, the others being metacentric elements. Zaedyus pichiy has 62 elements, also with one-half being acrocentric. The three-banded armadillo, “bolita,” which can roll up into a complete ball, has the lowest chromosome number of all, with 2n = 38, and as expected, all of them are metacentric. Among armadillos, this is perhaps the most derived species. Finally, of the subterrestrial “mouse armadillos” (Chlamyphorus), only one specimen has ever been studied, Chlamyphorus truncates; it had 58 chromosomes. They are never kept successfully in captivity and are only very rarely seen. Kurt Benirschke

ORDER XENARTHRA

Choloepus didactylus (Two-Toed Tree Sloth)

2n=53

Nash (unpublished)

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Choloepus hoffmanni (Two-Toed Tree Sloth/Hoffmann’s Sloth)

2n=50

Jorge et al. (1977) Contributed by K. Benirschke

ORDER XENARTHRA

Myrmecophaga tridactyla (Giant Anteater)

2n=60

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

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Tamandua tetradactyla (longicaudata) (Lesser Anteater/Tamandua)

2n=54

Jorge et al. (1977) Contributed by K. Benirschke

ORDER XENARTHRA

Cyclopes didactylus (Silky Anteater)

2n=64

Jorge (2000)

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Chaetophractus villosus (Hairy Armadillo/Peludos)

2n=60

Jorge et al. (1977) Contributed by K. Benirschke

ORDER XENARTHRA

Euphractus sexcinctus flavimanus (Six-Banded Armadillo)

2n=58

Jorge et al. (1977) Contributed by K. Benirschke

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Zaedyus pichiy (Pichi)

2n=62

Jorge et al. (1977) Contributed by K. Benirschke

ORDER XENARTHRA

Cabassous centralis (Naked-Tailed Armadillo)

2n=62

Benirschke et al. (1969)

Cabassous tatouay (Naked-Tailed Armadillo)

2n=50

Benirschke et al. (1969)

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Tolypeutes matacus (La Plata Three-Banded Armadillo)

2n=38

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Dasypus novemcinctus (Long-Nosed Armadillo/Nine-Banded Armadillo)

2n=64

Jorge et al. (1977) Contributed by K. Benirschke

ORDER XENARTHRA

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Dasypus septemcinctus (Long-Nosed Armadillo/Seven-Banded Armadillo)

2n=64

Barroso and Seuanez (1991) Contributed by H. Seuanez

Dasypus hybridus (Long-Nosed Armadillo)

2n=64

Jorge et al. (1977) Contributed by K. Benirschke

EUARCHONTOGLIRES Order Scandentia The tree shrews, classified in the order Scandentia, are a group of smallbodied, arboreal, insectivorous mammals endemic to south and southeastern Asia (from the Indian subcontinent and southern China to the Philippines, Borneo, and Bali). The higher level relationships of tree shrews have long been controversial and have attracted considerable study as a result. Traditionally, tree shrews have been classified either within the order “Insectivora” (an inclusive grouping of small-bodied, plesiomorphic placental mammals now known to be polyphyletic) or as basal members of the order Primates. Tree shrews are now known to be most closely related to colugos (order Dermoptera) and primates (Murphy et al., 2001b) but are best classified as a distinct order, Scandentia (Luckett, 1980). Current classifications recognize about 20 species and 5 genera of tree shrews (Olson et al., 2004; Helgen, 2005). Traditionally, these genera have been united in a single family, Tupaiidae, but recent assessments divide the order into two distinctive families—Tupaiidae (the true tree shrews, including Tupaia, Urogale, Anathana, and Dendrogale) and Ptilocercidae (the feather-tailed tree shrew, with the single genus Ptilocercus) (Shoshani and McKenna, 1998; Helgen, 2005). Together, the papers of Borgaonkar (1969), Arrighi et al. (1969), and Toder et al. (1992) admirably summarize current knowledge of tree shrew cytogenetics. In total, only two of the five currently recognized tree shrew genera (Tupaia and Urogale) have been studied cytogenetically. In Tupaia, six species have been karyotyped. Though the diploid complement varies from 2n = 52 (T. palawanensis) to 2n = 68 (T. montana), with most species having a relatively high diploid number (T. glis, 2n = 60; T. longipes, 2n = 60; T. belangeri, 2n = 62; T. minor, 2n = 66), the fundamental number is generally stable across the genus, at 72 (Borgaonkar, 1969). These chromosomal complements are described in some detail by Arrighi et al. (1969) and have helped in certain cases to distinguish between closely related tree shrew species. For example, Tupaia glis, T. belangeri, and T. palawanensis were often considered conspecific before these karyological studies, but their distinctness has since been confirmed on the basis of morphological and molecular data as well (e.g., Endo et al., 2000; Han et al., 2000). In other cytogenetic studies involving the genus Tupaia, Toder et al. (1992) described the number and position of the nucleolar organizing regions (NORs) in T. glis and T. belangeri, and Müller et al. (1999) discussed the evolution of tupaiid karyotypes in the context of a multidirectional chromosomal painting study of tree shrews and primates. Atlas of Mammalian Chromosomes, Edited by Stephen J. O’Brien, Joan C. Menninger, William G. Nash Copyright © 2006 John Wiley & Sons, Inc.

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EUARCHONTOGLIRES In contrast to Tupaia, the published karyotype of the monotypic Philippine genus Urogale everetti reveals a diploid number of 2n = 44 and an apparent fundamental number (FN) of 84 (Chu and Bender, 1962; Borgaonkar, 1969; Arrighi et al., 1969). This suggests that karyological data may be relevant to investigations regarding the boundaries and interrelationships of tupaiid genera, some aspects of which are still controversial (e.g., the monophyly of Tupaia with respect to Urogale and Anathana). In this light, karyotypic data for the unstudied genera Anathana and Dendrogale, and especially for Ptilocercus, the most basal living tree shrew (Sargis, 2004), would likely complement ongoing morphological and molecular studies of evolution within this distinctive order. Kristofer M. Helgen

ORDER SCANDENTIA

Tupaia belangeri (Northern Tree Shrew)

2n=62

Bigoni and Stanyon (unpublished)

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Order Dermoptera The order Dermoptera includes a single family, Cynocephalidae, with two living species: Galeopterus variegates, occurring in southern Indochina, the Malay Peninsula, Borneo, and other Sundaic islands (Corbet and Hill, 1992), and Cynocephalus volans, restricted to the Mindanao faunal region of the southern Philippines (Heaney et al., 1998). Known as flying lemurs, colugos, or kagwang, dermopterans are medium-sized (1–2-kg) gliding mammals that are nocturnal and arboreal and feed almost exclusively on leaves. They are characterized by fur-lined gliding membranes, simple molars, and specialized comblike lower incisors (Lekagul and McNeely, 1998; Stafford and Szalay, 2000; Wischusen and Richmond, 1998). Fossil dermopterans are known from the late Eocene of Thailand (Ducrocq et al., 1992). The extinct families Paromomyidae, Plagiomenidae, and Mixodectidae from the Paleocene and Eocene of North America have been variously considered to be either dermopterans or primates (Carroll, 1988; Stafford and Szalay, 2000). The relationship of Dermoptera to other mammalian orders has been a subject of some interest and contention (Stafford and Szalay, 2000). Dermopterans have been grouped in the grandorder Archonta along with Chiroptera, Primates, and Scandentia (Gregory, 1910; McKenna and Bell, 1997). On the basis of morphology, they have been considered the sister group of bats (Novacek and Wyss, 1986; Simmons, 1995) or more closely related to primates (McKenna and Bell, 1997). Molecular data unite Dermoptera, Scandentia, and Primates as a monophyletic group separate from Chiroptera (Adkins and Honeycutt, 1991; Schmitz et al., 2002). Most recent authors place the two extant dermopteran species in a single genus, Cynocephalus (Corbet and Hill, 1992; Wilson, 1993). However, based on the magnitude of morphological differences, Stafford and Szalay (2000) place them in separate genera (Cynocephalus volans and Galeopterus variegatus). Substantial chromosomal differences also support this separation. The karyotype of G. variegatus (Hsu and Benirschke, 1973) is 2n = 56, consisting of 20 pairs of telocentric and 7 pairs of biarmed autosomes, a large metacentric X, and a minute telocentric Y chromosome. The smallest autosome has a secondary constriction near the centromere. The karyotype of C. volans (Rickart, 2003) is 2n = 38, consisting of 18 pairs of small- to large-sized telocentric autosomes and a medium-sized submetacentric X chromosome (the Y chromosome is unknown). Distinct Gbands are present on the 9 largest autosomes. The smaller autosomes and X chromosome are largely G-positive. Silver-stained nucleolus organizer regions (Ag-NORs) are located terminally on the smallest telocentric autosomes. Eric A. Rickart

ORDER DERMOPTERA

Cynocephalus volans (Colugo/Flying Lemur/Kagwang)

2n=38

Rickart (2003)

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Order Primates The primates are a widespread order of placental mammals. Most species have a tropical or subtropical distribution and very few species venture into temperate zones. The primates can be divided into prosimians, New World primates (Platyrrhini), and Old World primates (Catarrhini). Catarrhini includes Old World monkeys, lesser ape, great apes, and our own species. The cytogenetics of primates can be logically taken back to 1956 when Tjio and Levan finally established the correct human diploid number (2n = 46). It is an interesting scientific footnote that the correct diploid number of the chimpanzee (2n = 48) was published 16 years earlier (Yeager et al., 1940). However, the study of primate chromosomes began in earnest in the early 1960s with the reports of Chiarelli (1962a,b; 1963; summarized in Chiarelli, 1968), Chu and Bender (1962), Hamerton et al. (1963), and Wohnus and Benirschke (1966). These early explorers, working with simple staining techniques, established the diploid and fundamental number (number of chromosome arms) of many species. In the 1970s chromosome-banding techniques were developed. The most widely used method was G-banding, but a distinctive and prolific French-inspired school developed which relied on Rbanding. Foremost among these workers was B. Dutrillaux (cf. Dutrillaux, 1979, for a review of his early results in the primates and a special supplement of Mammalia, vol. 50, 1986). In the late 1980s molecular cytogenetic work began. Comparative chromosome painting of nonhuman karyotypes sometimes dubbed Zoo-FISH (fluorescence in situ hybridization) has been the most important molecular cytogenetic technique. Chromosome paints employ DNA probes specific for single chromosomes. Human paints are now made by fluorescence-activated cell sorting (FACS), and this technique has also been used to make chromosome probe sets for nonhuman primates. This technique allows us to visualize one type of chromosome rearrangement, translocations. Inversion, deletions, and amplifications are best documented with other methods. However, chromosome painting is the only molecular method for which we have reasonable samples across the primate order. The brief review of primate chromosome here relies on data from all these techniques.

Great Apes 2n = 48 From the earliest years of cyotogenetics it was recognized that the chromosomal appearance of all great apes’ chromosomes was amazingly similar to humans. We know now that the vast majority of human syntenies are found intact in these species. As expected the synteny of human chromosome 2 was not conserved. Two pairs of homologues are found in all species because human chromosome 2 is the result of a recent fusion in the evolutionary line leading to humans. This accounts for the difference in diploid number between human (2n = 46) and all great apes (2n = 48). The only other synteny alteration was a derived (apomorphic) reciprocal translocation between homologues to human chromosome 5 and 17 in the gorilla.

Lesser Apes 2n = 38, 44, 50, 52 Hylobatids (gibbons, or lesser apes) are classified with great apes and human in the same primate superfamily Hominoidea; however, the origin, phylogeny,

ORDER PRIMATES and homology of their chromosomes were never identified by chromosomebanding studies. In fact, because chromosomal homologies between lesser apes and other primates including human were so difficult to propose solely on the basis of banding, they were systematically left out of chromosomal phylogenies of the primates. Recent literature identifies about 12 distinct species. Chromosome studies showed four main karyotypes with a wide range of diploid chromosome numbers (Hylobates hoolock: 2n = 38, H. lar species group: 2n = 44, H. concolor: 2n = 50, H. syndactylus: 2n = 52). Chromosomal painting revealed that synteny relative to human and great apes was highly disturbed. It is clear that lesser apes have experienced rapid and massive chromosome evolution and as a consequence had highly derived karyotypes marked by many translocations. For instance, there has been a minimum of 33 translocations in the evolution of the siamang karyotype. The 24 autosomes are composed of 60 recognizable segments that show DNA homology to regions of the 22 human autosomes. There are also profound differences among lesser apes. For example, since the phylogenetic separation of H. lar and H. syndactylus these two lesser apes have independently accumulated numerous translocations (16 in H. syndactylus and 14 in H. lar). Thus, the two hylobatids are separated from each other by a total of 30 translocations and numerous intrachromosomal rearrangements. Gibbons are unique among higher primates in showing extensive chromosome polymorphisms, which include not only inversions but also translocations (Couturier et al., 1982; van Tuinen and Ledbetter, 1983; Stanyon et al., 1983, 1987b; Couturier and Lernould, 1991). However, polymorphisms for inversions can also be observed within and between subspecies of the orangutan. Even though very few gibbons have been karyotyped, all species have chromosome polymorphisms suggesting that the process of karyological transformation is still underway. Chromosome painting identified inversions, a transposition, and a translocation polymorphism. Apparently, all H. lar species group gibbons analyzed share an inversion polymorphism on chromosome 8 (Stanyon et al., 1987a), which indicates that the polymorphism is not transient but has even survived speciation events. However, for some heterozygote rearrangements it is not totally clear whether they arose from breeding animals from different populations or subspecies. This has to be further analyzed, including larger sample sizes with animals of known geographic origin and preferentially free-ranging individuals. The reason why hylobatids are so chromosomally derived remains elusive. It is not clear whether gibbons have a higher chromosomal mutation rate or the mutations which occur are simply more easily fixed or both. A rapid fixation may have been favored by gibbon social structure and ecology, including monogamous matings, nuclear family units, and an arboreal lifestyle. In contrast, the chromosomally conservative Papionini (see below) live in large terrestrial groups with multiple male and multiple female matings. Population bottlenecks and inbreeding also have been proposed to explain the rapid fixation of such rearrangements, but the extensive chromosome polymorphism which is even shared by different species would argue against drastic bottlenecks during the divergence of gibbons. However, the results clearly demonstrate that a “molecular clock” as proposed for gene mutations over time of evolution would not hold true for chromosome rearrangements since, as will be shown in more detail, even more distantly related species than gibbons (various Old World monkeys) have much more conserved karyotypes compared to human.

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Old World Monkeys 2n = 42–72 Old World monkeys are divided into two families: Cercopithecidae (including baboons, macaques, guenons, etc.) and Colobidae (African and Asian leafeating monkeys). Only a few translocations have taken place in the evolution of these species, but the guenons and their allies (cercopithecini) are characterized by chromosome fissioning (accounting for all diploid numbers above 46 found in these monkeys). The papionini, which includes macaques, baboons, mangabeys, drills, and mandrills all have nearly identical karyotypes. All human chromosome syntenies except for the homologue for human chromosome 2 are found intact. However, three chromosomes are composed of two human chromosomes: 7/21, 14/15, and 20/22. These associations between different human chromosomes account for the diploid number of 2n = 42 found in these monkeys. In Colobinae, the other major division of Old World monkeys, the diploid number of 2n = 44 is closer to that of humans, but more translocation and resulting associations, a combination of human syntenies, were found. In the black and white colobus (Colobus guereza) the 24 human paints gave 31 signals on the autosomes (haploid chromosome set). Reciprocal translocations were found between human chromosomes 1/10, 1/17, as well as 3/19. A different set of translocation was found in the silvered leaf monkey (Trachypithecus cristatus): between 1/19, 6/16, and 5/Y. The proboscis monkey is the only species of Colobinae with a different diploid number, 2n = 48. This higher diploid number is explained by derived fissions of a segment of human chromosomes 14 and 6.

New World Monkeys 2n = 16–62 Cytogenetic studies have shown that New World primates are karyologically diverse and some species are highly derived with respect to the chromosome complement of man or a presumed platyrrhine ancestor. Current taxonomy classifies the New World monkeys (platyrrhini) into two families, Cebidae and Callithrixidae. There is now ample painting data on chromosome translocations and synteny associations from many species in each family. Previously some scientists proposed that these monkeys could have had two distinct (polyphyletic) origins perhaps independently from North America and Africa. However, there are a number of translocation associations of homologues to human chromosomes (8/18, 10/16, 2/16, 5/7), which unite both families of New World monkeys and prove without a doubt that they are monophyletic (had one common origin). Most primatologists now think that these monkeys had an African origin. The taxonomy and phylogeny of neotropical primates are still debated. Recently molecular cytogenetic data on translocations present in these species helped clarify the relationship of Goeldi’s marmoset (Callimico goeldii). The taxonomic assessment of this species has varied from author to author since it was discovered about 100 years ago. Some primatologists even assigned it to its own family. However, Callimico species share chromosome rearrangements. The analysis of the chromosome painting data clearly indicates that Callimico shares numerous translocations and associations (16/19, 2/15, 9/13, 9/22, 13/17, 17/20) in common with and is nested within Callitrichidae. Comparisons with Saimiri sciureus show that this species exclusively shares a derived syntenic association of human 2/15 homologues with all Callithrichidae, suggesting that this genus is a sister group.

ORDER PRIMATES Comparative cytogenetic research has also shown that New World monkey biodiversity is not yet well known and that the number of species might be underestimated by traditional taxonomy. The recent finding of a new species of Callicebus with the lowest diploid number in all of the primate order 2n = 16 illustrates this well.

Prosimians 2n = 36–80 Prosimians are a highly diversified catch-all group that includes lemurs, lorids, and tarsiers. Lemurs are exclusively found on the island of Madagascar and the R-banding karyotypes of about 17 species have been described. Comparisons of karyotypes between lemur species are not too difficult to interpret since apparently they mostly consist of Robertsonian chromosome fusions or fissions. Some lemur karyotypes have unusual features such as microchromosomes which even in high-resolution metaphase preparations lack distinct banding patterns. Among the lemurs Cheirogalaeidae has a diploid number of 66 with the exception of the genera Phaner (2n = 48). The karyotype of Microcebus 2n = 66 is often been selected as ancestral to lemurs and other primates. The prosimians (lemurs and lorids), in general, are often assumed to be more primitive and therefore more similar to the ancestor of all primates than monkeys, apes, and humans. Indeed, on the basis of chromosome banding, some authors have suggested that the chromosomes of some prosimian species were close to the ancestral primate karyotype. Painting data do not support the conclusions of the chromosome-banding studies; in fact, the number of apomorphic chromosomal syntenies found in both Lemuridae and Lorisidae demonstrates that these species are not close to the assumed ancestral primate karyotype. The phylogenetic position and taxonomy of the genus Tarsius is highly debated. Some taxonomists place this species on the line leading to higher primates, while others place it among the prosimians. Diploid numbers among Tarsius vary from the highest number found in primate, 2n = 80, with some species with diploid number of 66 and 46. Cytogenetic studies of the tarsier are rare and not informative about the phylogenetic position of the tarsier.

Ancestral Primate Karyotypes Several attempts to construct the ancestral primate chromosome organization have appeared (Haig, 1999; Chowdhary et al., 1998; Müller et al., 1999; O’Brien and Stanyon, 1999 and Ruiz-Herrera et al., 2005). The associations 4/8p, 7/16, and 16/19 present in the ancestral placental mammal karyotype are broken up in the prosimians and two chromosomes are formed. Comparison suggests that chromosomes 4, 8, 16a, 16b, 19a, and 19b could be included into the ancestral primate karyotype. Fusions would have given origin to the syntenies of 7, 8, and 10. The ancestral primate karyotype would then include the chromosome forms 1a, 1b, 2a, 2b, 3/21, 4–11, 12a/22a, 12b/22b, 13, 14/15, 16a, 16b, 17, 18, 19a, 19b, 20, and X and Y and would have had a diploid chromosome number of 2n = 50. Of these forms, chromosomes 1a, 1b, 4, 8, 12a/22a, and 12b/22b are probably common derived characters that would link the tree shrew with primates. Anthropoid origins would be marked by a reciprocal translocation to form chromosomes homologous to 12 and 22 and a fusion gave origin to chromosome 19 yielding a diploid number of 48. New World monkeys are all linked by fragmentation of chromosomes 1, 2, 7, 8, 10, and 15 as well as translocations to form associations 2/16, 5/7, 8/18,

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EUARCHONTOGLIRES and 10/16. Consequently, the ancestral New World monkey karyotype should have had a diploid number of 2n = 54 chromosomes. It consisted of homologues to human chromosomes 1a, 1b, 1c, 2a, 2b/16b, 3a/21, 3b, 3c, 4, 5/7a, 6, 7b, 8a/18, 8b, 9, 10a/16a, 10b, 11, 12, 13, 14/15a, 15b, 17, 19, 20, 22, X and Y, this assuming that the ancestral karyotype is found conserved in the genus Cebus. The evolution of the ancestral catarrhine karyotype would have involved the fusion of chromosomes 1 and 16 and the fission of the ancestral synteny between 3 and 21. The chromosome 14/15 association, which is found in many outgroups of nonprimates, would also be present in the ancestral catarrhine karyotype. The diploid number of the ancestral catarrhine karyotype would then be 2n = 46, including homologues to human chromosomes 1, 2a, 2b, 3–13, 14/15, 16–22, X, and Y. All hominoids (sl) are linked by fissions of ancestral mammalian synteny 14/15. Roscoe Stanyon

ORDER PRIMATES

Loris tardigradus (Slender Loris)

2n=62

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

105

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Nycticebus pygmaeus (Pygmy Slow Loris/Cu Lan)

2n=50

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER PRIMATES

Nycticebus coucang (Slow Loris/Cu Lan)

2n=50

Stanyon et al. (1987a)

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Galago moholi (Galago/Bushbaby)

2n=38

Stanyon et al. (2002)

ORDER PRIMATES

Otolemur crassicaudatus (Brown Galago)

2n=62

Stanyon et al. (2002)

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Microcebus murinus (Mouse Lemur)

2n=66

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Eulemur macaco flavifrons (Black Lemur)

2n=44

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER PRIMATES

Eulemur fulvus sanfordi (Sanford’s Lemur)

2n=60

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

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Lepilemur ruficaudatus (Red-Tailed Sportive Lemur )

R-bands 2n=20

Rumpler et al. (1985)

Lepilemur Ieucopus (White-Footed Sportive Lemur)

R-bands 2n=26

Rumpler et al. (1985)

ORDER PRIMATES

Lepilemur septentrionalis (Northern Sportive Lemur)

R-bands 2n=34

Rumpler et al. (1985)

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Propithecus verreauxi coquereli (Coquerel’s Sifaka)

2n=48

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Propithecus tattersalli (Golden-Crowned Sifaka)

2n=42

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER PRIMATES

Daubentonia madagascariensis (Aye-Aye)

2n=30

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Ideogram: Daubentonia madagascariensis (Aye-Aye)

2n=30

Tagle et al. (1990) Contributed by D. A. Miller

* Q-Bands

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Tarsius bancanus (Western Tarsier)

2n=80

Poorman et al. (1985)

ORDER PRIMATES

Tarsius syrichta (Philippine Tarsier)

2n=80

Rickart (2003)

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Lagothrix Iagotricha (Woolly Monkey)

2n=62

Stanyon et al. (2001)

2n=44(F)

Stanyon et al. (1995)

Alouatta seniculus (Red Howler)

ORDER PRIMATES

Alouatta sara (Bolivian Red Howler)

2n=52

Stanyon et al. (1995)

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Alouatta belzebul (Red-Handed Howler Monkey)

2n=50(F) 49(M)

Consigliere et al. (1998) Contributed by R. Stanyon *(Y translocated to autosome)

ORDER PRIMATES

Ideogram: Alouatta belzebul (Red-Handed Howler Monkey)

2n=50(F) 49(M)

Consigliere et al. (1998) Contributed by R. Stanyon

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Alouatta caraya (Black Howler Monkey)

2n=50

Bigoni and Stanyon (unpublished)

ORDER PRIMATES

Pithecia pithecia (White-Faced Saki)

2n=48

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

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Cacajao calvus rubicundus (Red Uakari/Uacari)

2n=46

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER PRIMATES

Callicebus cupreus (Coppery Titi Monkey)

2n=46

Bigoni and Stanyon (unpublished)

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Callicebus pallescens (White-Coated Titi Monkey)

2n=50

Stanyon et al. (2000)

Callicebus personatus nigrifrous (Atlantis Titi Monkey)

2n=42

Nagamachi et al. (2003)

ORDER PRIMATES

Aotus nancymaae (Nancy Ma’s Night Monkey/Douroucouli)

2n=54

F. Garcia et al. (2002b)

Aotus azarae (Azara’s Night Monkey/Douroucouli)

Contributed by M. Ponsa

2n=50

F. Garcia et al. (2002b) Contributed by M. Ponsa

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Cebus capucinus (White-Headed Capuchin Monkey/ Azara’s White-Faced Monkey/Sapajon)

2n=54

Campa and Stanyon (1992) Contributed by R. Stanyon

ORDER PRIMATES

Cebus albifrons albifrons (White-Fronted Capuchin Monkey)

2n=54

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

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Cebus apella (Tufted Capuchin Monkey)

2n=54

Clemente et al. (1987) Contributed by M. Ponsa

Ideogram: Cebus apella (Tufted Capuchin Monkey)

2n=54

Clemente et al. (1987) Contributed by M. Ponsa

ORDER PRIMATES

Cebus olivaceus (Weeper Capuchin)

2n=52

F. Garcia et al. (2002b) Contributed by M. Ponsa

2n=44

Stanyon et al. (2000)

Saimiri sciureus (Squirrel Monkey)

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Callimico goeldii (Goeldi’s Marmoset)

2n=48(F)

47(M)

Bigoni and Stanyon (unpublished)

Saguinus imperator (Emperor Tamarin)

2n=46

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER PRIMATES

Saguinus midas (Red-Handed Tamarin)

2n=46

Bigoni and Stanyon (unpublished)

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Callithrix jacchus (Common Marmoset)

2n=46

Bigoni and Stanyon (unpublished)

Callithrix argentata (Silvery Marmoset)

2n=44

Canavez et al. (1996) Contributed by H. N. Seuanez

ORDER PRIMATES

Callithrix emiliae (Emilia’s Marmoset)

2n=44

Nagamachi et al. (1996)

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Callithrix humeralifer (Santarem Marmoset)

2n=44

Nagamachi et al. (1996)

ORDER PRIMATES

Callithrix chrysoleuca (Gold and White Marmoset)

2n=44

Nagamachi et al. (1996)

Callithrix mauesi (Maues Marmoset)

2n=44

Nagamachi et al. (1996)

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Cebuella pygmaea (Pygmy Marmoset)

2n=44

Bigoni and Stanyon (unpublished)

ORDER PRIMATES

Leontopithecus chrysopygus (Black Lion Tamarin)

2n=46

Seuanez et al. (1988)

Leontopithecus rosalia (Golden Lion Tamarin)

2n=46

Seuanez et al. (1988)

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Erythrocebus patas (Patas Monkey)

2n=54

Bigoni and Stanyon (unpublished)

ORDER PRIMATES

Cercopithecus (Chlorocebus) aethiops (African Green Monkey/Grivet Monkey)

2n=60

Bigoni and Stanyon (unpublished)

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Ideogram: Cercopithecus (Chlorocebus) aethiops (African Green Monkey/Grivet Monkey)

2n=60

Finelli et al. (1999) Contributed by R. Stanyon

ORDER PRIMATES

Cercopithecus nictitans (Greater Spot-Nosed Guenon/ White-Nosed Monkey)

2n=70

Ponsa and Eqozcue (1981)

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Cercopithecus ascanius schmidti (Schmidt’s Spot-Nosed Guenon/ Red-Tailed Monkey)

2n=66

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER PRIMATES

Cercopithecus petaurista (Lesser Spot-Nosed Guenon)

2n=66

Clemente et al. (1990) Contributed by M. Ponsa

Cercopithecus mona campbell (Mona Monkey)

2n=66

Clemente et al. (1990) Contributed by M. Ponsa

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Cercopithecus cephus (Mustached Guenon)

2n=66

Clemente et al. (1990) Contributed by M. Ponsa

ORDER PRIMATES

Cercopithecus diana (Diana Monkey)

2n=58

Ponsa et al. (1986)

2n=72

Ponsa and Eqozcue (1981)

Cercopithecus pogonias (Guenon)

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Cercopithecus I’hoesti (L’Hoest Monkey/L’Hoest Guenon)

2n=60

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER PRIMATES

Miopithecus talapoin (Talapoin)

2n=54

Ponsa et al. (1980)

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Allenopithecus nigroviridis (Allen’s Swamp Monkey)

2n=48

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER PRIMATES

Cercocebus galeritus (Tana River Mangabey)

2n=42

Bigoni and Stanyon (unpublished)

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Cerocebus torquatus (Collared Mangabey/Red-Capped Mangabey)

2n=42

F. Garcia et al. (2002) Contributed by M. Ponsa

2n=42

Ruiz-Herrera et al. (2002) Contributed by M. Ponsa

Macaca fascicularis (Crab-Eating Macaque)

ORDER PRIMATES

Macaca mulatta (Rhesus Monkey)

2n=42

Bigoni and Stanyon (unpublished)

Papio hamadryas (Hamadryas Baboon)

2n=42

Bigoni and Stanyon (unpublished)

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Macaca fuscata (Japanese Macaque)

2n=42

Bigoni and Stanyon (unpublished)

ORDER PRIMATES

Macaca fuscata (Japanese Macaque)

2n=42

Wienberg et al. (1992) Contributed by R. Stanyon

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Mandrillus sphinx (Mandrill)

2n=42

Ponsa, F. Garcia, M. Garcia, Ruiz-Herrera, and Eqozcue (unpublished)

Trachypithecus obscurus (Dusty Leaf Monkey)

2n=44

Ponsa et al. (1983)

ORDER PRIMATES

Pygathrix nemaeus (Douc Langur)

2n=44

Bigoni (1995)

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Trachypithecus cristatus (Silver Leaf Monkey)

2n=44(F)

44(M)

Bigoni et al. (1997a)

ORDER PRIMATES

Colobus guereza (Black and White Colobus Monkey/Kikuya)

2n=44

Bigoni et al. (1997b)

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160

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Hylobates syndactylus (Siamang)

2n=50

Ponsa, F. Garcia, M. Garcia, Ruiz-Herrera, and Eqozcue (unpublished)

Hylobates concolor (Black-Crested Gibbon/Concolor Gibbon)

2n=52

Bigoni and Stanyon (unpublished)

ORDER PRIMATES

Ideogram: Hylobates concolor (Black-Crested Gibbon/Concolor Gibbon)

2n=52

Koehler et al. (1995) Contributed by R. Stanyon

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Ideogram: Hylobates lar (White-Handed Gibbon)

2n=44

Jauch et al. (1992) Contributed by R. Stanyon

ORDER PRIMATES

Ideogram: Hylobates moloch (Silvery Gibbon)

2n=42

Stanyon and Chiarelli (1983)

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Pongo pygmaeus abelii (Sumatran Orangutan/Orangutan)

2n=48

Bigoni and Stanyon (unpublished)

ORDER PRIMATES

Ideogram: Pongo pygmaeus (Sumatran Orangutan/Orangutan)

2n=48

Yunis and Prakash (1982)

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Gorilla gorilla (Lowland Gorilla)

2n=48

Bigoni and Stanyon (unpublished)

ORDER PRIMATES

Ideogram: Gorilla gorilla (Lowland Gorilla)

2n=48

Yunis and Prakash (1982)

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Pan troglodytes (Common Chimpanzee)

2n=48

Bigoni and Stanyon (unpublished)

ORDER PRIMATES

Ideogram: Pan troglodytes (Common Chimpanzee)

2n=48

Yunis and Prakash (1982)

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Pan paniscus (Pygmy Chimpanzee/Bonobo)

2n=48

Bigoni and Stanyon (unpublished)

ORDER PRIMATES

Homo sapiens (Human)

2n=46

Padilla-Nash (unpublished)

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Ideogram: Homo sapiens (Human)

Francke (1994)

* nontraditional format due to size limitations

ORDER RODENTIA

Order Rodentia Rodentia is the largest order of mammals encompassing at least 40% of recognized mammalian species. Rodents are ubiquitous and occur in all continents except Antarctica. Most species are small sized and they are usually vegetarian. Rodents cost billions of dollars in crop losses each year. Some are carriers of human diseases such as bubonic plague, typhus, and hantaviral disease. However, various rodent species are economically important as food source and in the fur industry and others are used extensively in biomedical research. The subordinal division of rodents is not yet clear. In the light of recent molecular studies neither Brandt’s (1855) classification recognizing three suborders (Sciuromorpha, Myomorpha, and Hystricomorpha), nor that of Tullberg (1899) recognizing two (Sciurognathi and Hystricognathi) are completely satisfactory. Waiting for the new taxonomic revision integrating molecular data, here we will follow the classification of Tillberg, which is currently accepted (Musser and Carleton, 1993). The development of rodent cytogenetics is associated first of all with the name of Robert Matthey, who, since the 1940s, has applied systematic chromosomal analysis to the study of vertebrate taxonomy, an approach he called cytotaxonomy. This alpha taxonomy was completed with the publication of the Atlas of Mammalian Chromosomes by T. C. Hsu and K. Benirschke. The wide application of banding techniques in mammalian cytogenetics was initiated in Hsu’s laboratory.

Chromosome Numbers (For detailed description of chromosomal sets see: www.bionet.nsc.ru\ chromosomes.) The suborder Sciurognathi includes 2021 species classified in 443 genera belonging to 11 families (Musser and Carleton, 1993). The mountain beaver, Aplodontia rufa, from the monospecific family Aplodontidae, has 2n = 46. The family Sciuridae (squirrels, ground squirrels, marmots, chipmunks) contains 273 species in 50 genera with chromosomal numbers varying from 2n = 30 in some species of the genus Citellus to 2n = 62 in Menetes berdmorei. Two species of the family Castoridae (beavers), Castor fiber and C. canadensis, possess 2n = 48 and 2n = 40, respectively. Within the family Geomyidae (pocket gophers) with 35 species in 5 genera, the diploid numbers range between 38 in Geomys tropicalis and 82 in Thomomys bottae pervagus. The family Heteromyidae (pocket mice, kangaroo mice, kangaroo rats, and spiny pocket mice) is formed by 59 species distributed into 6 genera. Chromosome numbers vary from 2n = 34 in Perognathus hispidus to 2n = 74 found in some species of the genus Dipodomys. Chromosome numbers in the species of the family Dipodidae (jumping mice) range between 32 in Sicista betulina and 72 in Zapus hudsonius. In Dipodidae (jerboas), chromosome numbers vary from 2n = 46 in Salpingotus crassicauda to 2n = 58 in Stilodipus telum. The most speciose family Muridae is formed by 1326 species distributed in 281 genera and 17 subfamilies. Among the species of the subfamily Murinae (Old World rats and mice) diploid numbers range between 16 in Mus (Leggada) goundae to 76 found in Grammomys macmillani (earlier as G. gazellae). In the New World rats and mice of the subfamily Sigmodontinae, the second largest subfamily, diploid numbers vary from 2n = 9, 10 in Akodon sp. to 2n = 92 in Anotomys leander and Ichthyomys pittieri. Chromosomal variation in Arvicolinae (voles) ranges from 2n = 17 in Ellobius lutescens and Microtus oregoni to 2n = 62 in some species of the genera Pitymys and Microtus. In the species of the subfamily

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EUARCHONTOGLIRES Cricetinae (hamsters) diploid numbers range between 20 in some species of the genus Cricetulus and 52 in some species of the genus Calomyscus. Chromosome numbers in Gerbillinae vary from 2n = 14, 15 in Taterillus tranieri to 2n = 74 in some species of the genus Gerbillus. The family Anomaluridae (scaly-tailed squirrels) is formed by seven species and three genera, showing diploid numbers ranging between 38 and 48. The monotypic Pedetidae (African springhare) includes Pedetes capensis having 2n = 38. In two species of the genus Ctenodactylus (Ctenodactylidae) karyotypes are similar and have 2n = 48. Among species of the family Myoxidae (dormice) diploid numbers vary from 44 in Myomimus personatus to 62 in Myoxus glis. The suborder Histricognathi includes 18 families but only 68 genera and 229 species. Most species of this suborder are large-sized rodents distributed in the Neotropics and Africa. In the species of the family Bathyergidae (African mole-rats) diploid numbers range between 2n = 40 in Cryptomys mechowi and 2n = 60 in Heliophobius argenteocinereus and Heterocephalus glaber. In Hystricidae (Old World porcupines), 2n varies from 46 in Trichys fasciculata (formerly as T. lipura) to 66 in some Hystrix species; in Thryonomyidae (cane rats) Thryonomys swinderianus has 2n = 44; in the family Erethizontidae (New World porcupines), including 4 genera and 12 species, the only karyotype reported is that of Erethizon dorsatum, having 2n = 42. In the family Chinchillidae (chinchillas) Chinchilla laniger has 2n = 64 and the viscacha Lagostomus maximus has 2n = 58. The chromosomally conservative family Caviidae (Guinea pigs) is formed by 14 species and 5 genera showing diploid number of 2n = 64 in all karyotyped species of the genus Cavia and 2n = 68 in Galea musteloides. The monotypic Hydrochaeridae contains the capybara Hydrochaeris hydrochaeris, the world’s largest rodent, having 2n = 66. In the family Agoutidae (pacas) diploid numbers vary from 2n = 42 in Agouti paca to 2n = 74 in A. taczanowskii. Within the family of fossorial rodents Ctenomyidae (tuco-tucos) the diploid numbers range between 22 in Ctenomys occultus and 70 in Ctenomys sp. In the family Octodontidae (octodonts, or degus) the chromosome number is 38 in Octodontomys gliroides whereas in Tympanoctomys barrerae it is 102, the highest chromosome number within mammals (see below). Chromosomal variation in the specious family Echimyidae (American spiny rats), suborder Hystricognathi, is extensive, and it varies from 2n = 24 in some species of the genus Proechimys to 2n = 90 in Echimys sp. Two species of the family Capromyidae (hutias), namely Capromys pilorides and Geocapromys brownii, have 2n = 40 and 2n = 88, respectively. The coypu or nutria, Myocastor coypus, from the monospecific family Myocastoridae, possesses 2n = 42.

Chromosome Variability One of the most remarkable features of rodents is an extreme variability of their chromosome numbers and/or chromosome morphology that resulted from polymorphism for different kinds of structural rearrangements and/or variation of the heterochromatic component of their genomes. Often variation of diploid numbers within a species results from polymorphism for centric, or Robertsonian, translocations, sometimes involving all or most of the acrocentric chromosomes. Known as Robertsonian fusion, such a large variation was found in a row of species, in particular in Mus musculus, where all diploid numbers ranging between 22 and 40 occur. Taking into account that the order of fusions of acrocentrics into metacentrics may be different in different populations, the number of different karyotypes found in nature reaches 60 (Bauchau, 1990). Another Robertsonian fusion is revealed in Ellobius talpinus with 2n varying from 54 to 31 (Lyapunova et al., 1980). Twenty-four and 20 different karyotypes were found in Akodon cursor (Fagundes et al., 1997; Silva

ORDER RODENTIA and Yonenaga-Yassuda, 1998) and Gerbillus nigeriae (Volobouev et al., 1995), respectively. However in these species as well as in those from the genera Ctenomys and Thomomys, also displaying very large karyotypic variation, Robertsonian variability is superposed by other types of chromosomal changes. Probably the most important source of karyotype variability in rodents is related to heterochromatin amount and its distribution pattern detectable by C-banding techniques and directly correlated with nuclear DNA content. After the finding of the first example of karyotype evolution by entirely heterochromatic arm additions/deletions in the species of the genus Peromyscus (Pathak et al., 1973), this type of polymorphism was detected in numerous rodent species. Now it is well documented that heterochromatin content may vary greatly between karyotypes of congeneric species otherwise similar or different only by some structural rearrangements. The interspecies differences of heterochromatin amount expressed by quantity of nuclear DNA attain 33% in Dipodomys species (Hatch et al., 1976), 36% in Peromyscus species (Deaven et al., 1977), 42% in Ammospermophilus (Mascarello and Mazrimas, 1977), and 60% in Thomomys species, where DNA content ranges between 5.8 pg in T. monticola having very small blocks of C heterochromatin and 14.2 pg in T. bottae with very large heterochromatic blocks (Patton and Sherwood, 1982; Sherwood and Patton, 1982). Genome size in the spiny rat Proechimys is also large and associated with repetitive DNA sequences, as is the case of P. trinitatis (2n = 62), having 12.5 pg DNA (Garagna et al., 1997). Sometimes a rather important interspecific or even interindividual variation of C heterochromatin amount is limited exclusively to the sex chromosomes, which is the case with the species from the genera Nesokia (Juyal et al., 1989), Microtus (Modi, 1993; Modi et al., 2003), and Arvicanthis (Garagna et al., 1999). Polymorphism for supernumerary or B chromosomes (B’s) is a particular case of heterochromatin variation as most of the B’s do not contain coding DNA sequences. The B’s occur in about 1.5% of mammalian species, at least two-thirds of which are rodents, mostly of the family Muridae (Volobouev, 1981). Here we describe two spectacular examples of B’s variation, found in the collared lemming Dicrostonyx torquatus (= groenlandicus), 2n = 40 plus 1 to 42 B’s (Chernjavsky and Kozlovsky, 1980) and of Apodemus peninsulae with 2n = 48 plus 0 to 24 B’s, some of which may be larger than the largest A chromosomes (Volobouev, 1981).

Sex Chromosome Systems Although some cases of the sex chromosome systems other than XX/XY are found within various mammalian taxa, rodents present the largest range of chromosomal mechanisms of sex determination. Contrary to the belief that the Y chromosome is necessary to determine normal male development, some rodents do not have a Y chromosome (Graves, 2002; Just et al., 2002). Among these, the most intriguing case is found in the mole vole Ellobius lutescens with X0/X0 constitution in both sexes. All attempts to localize any of the Ylinked genes on the X chromosome were not successful (Just et al., 1995, 2002). It is worthy to note that embryonic mortality in this species resulting from elimination of the XX and 00 carriers is more than 50%. Both sexes of congeneric Ellobius talpinus and Ellobius tancrei possess isomorphous sex chromosomes with identical banding pattern, and one of these is late replicated in females as well in males. Although embryonic mortality is about 10%, the males are fertile (Fredga and Lyapunova, 1991). The Japanese ryukyu spiny rats Tokudaia osimensis osimensis also has no cytologically recognizable Y chromosome; the Y-linked functional genes, such as Tspy and Zfy, were found to be translocated onto the distal part of the long arm of the X chromosome (Honda et al., 1978; Arakawa et al., 2002).

175

176

EUARCHONTOGLIRES The wood lemming, Myopus schisticolor, is known to have a biased sex ratio due to the presence of fertile XY females. A modified X chromosome having a deletion in the short arm of the X and a complex rearrangement explains these findings (Fredga, 1988). Several species of South American Akodon also have XY females (Bianchi et al., 1993). There are numerous rodent species, mostly in the subfamily Gerbillinae, having XX/XY1Y2 sex chromosomes that resulted from X autosome translocation (Dobigny et al., 2002). In addition, X0 females were found in Microtus oregoni (Ohno et al., 1963, 1964) and Acomys selousi (Matthey, 1965).

Tetraploidization This form of chromosomal evolution is considered theoretically impossible in mammals, although seemingly not for rodents. However, the flow cytometry analysis of the red viscacha rat Tympanoctomys barrerae (Octodontidae) showed that size of the nuclear genome of this species is twice larger than that in relative species, 16.8 pg versus 8.7 pg (Gallardo et al., 1999, 2003). Tympanoctomys barrerae has the largest chromosome complement reported for a mammal, with 2n = 102, thus more probably being tetraploid. Its karyotype is formed by biarmed elements having prominent centromeric blocks of heterochromatin. The recently described octodontid species Pipanacoctomys aureus also has a double genome size and a totally biarmed karyotype with 92 chromosomes (Gallardo et al., 2004).

Comparative Genomics Until recently, only some rodent species profited from studies with modern molecular cytogenetics techniques. So chromosome painting of the mouse and comparative gene mapping of mouse and human revealed that mouse has the most rearranged karyotype of all mammals so far investigated (FergusonSmith et al., 1998). It turned out that the autosomal complement of M. musculus, consisting of 19 pairs, is divided into 180 syntenic segments (Genome Sequencing Consortium, 2001), and the 20-chromosome complement of the rat is distributed in more than 120 syntenic segments (Grutzner et al., 1999; Nilsson et al., 2001). The high level of fragmentation and repatterning of the mouse genome may probably account for the limited success of chromosome painting in rodents (Ferguson-Smith et al., 1998). In contrast, the karyotypes of two squirrel species, Sciurus carolinensis and Menetes berdmorei, are divided into 39 and 34 fragments, respectively. Squirrel genomes appear to be very similar to that of the rabbit, Oryctolagus cuniculus, as well as to that of the proposed mammalian ancestor (Richard et al., 2003; Stanyon et al., 2003; Li et al., 2004). Using M. musculus chromosome-specific probes, five species of Muridae (Mus platythrix, Rattus novegicus, Rattus rattus, Rhabdomys pumilio, and Cricetulus griseus) have been studied (Rabbitts et al., 1995; Grutzner et al., 1999; Guilly et al., 1999; Stanyon et al., 1999; Yang et al., 2000b; Cavagna et al., 2002; Matsubara et al., 2003; Rambau and Robinson, 2003). It follows from these analyses that the rate of chromosome translocations in rodents in approximately one order of magnitude higher than that between species from other mammalian orders, as human and cat. V. T. Volobouev Milton H. Gallardo Alexander S. Graphodatsky

ORDER RODENTIA

Tamias sibiricus (Chipmunk)

2n=38

Graphodatsky and Sablina (unpublished) Contributed by A. Graphodatsky

Sundasciurus philippinensis (Sunda Tree Squirrel)

2n=38

Rickart (2003)

177

178

EUARCHONTOGLIRES

Spermophilus erythrogenys (Ground Squirrel/Susliks) G-bands

2n=36

Graphodatsky and Sablina (unpublished) Contributed by A. Graphodatsky

C-bands

2n=36

Graphodatsky and Sablina (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Spermophilus undulatus (Ground Squirrel/Susliks) G-bands

2n=32

Graphodatsky and Sablina (unpublished) Contributed by A. Graphodatsky

C-bands

2n=32

Graphodatsky and Sablina (unpublished) Contributed by A. Graphodatsky

179

180

EUARCHONTOGLIRES

Spermophilus alaschanicus (Ground Squirrel/Susliks) G-bands

2n=38

Graphodatsky and Sablina (unpublished) Contributed by A. Graphodatsky

C-bands

2n=38

Graphodatsky and Sablina (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Petaurista albiventer (Chestnut Great Flying Squirrel)

2n=38

Li and Yang (unpublished) Contributed by F. Yang

181

182

EUARCHONTOGLIRES

Castor fiber (Beaver)

2n=48

Graphodatsky (unpublished)

ORDER RODENTIA

Castor canadensis (Beaver)

2n=40

Graphodatsky (unpublished)

183

184

EUARCHONTOGLIRES

Thomomys talpoides (Western Pocket Gopher)

2n=48

Thaeler (1974)

Thomomys talpoides attenuatus (Western Pocket Gopher)

2n=48

Thaeler (1968)

ORDER RODENTIA

Thomomys talpoides pygmaeus (Western Pocket Gopher)

2n=48

Thaeler (1968)

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186

EUARCHONTOGLIRES

G-bands R-bands DAPI C-bands

Ideogram: Geomys attwateri (Pocket Gopher)

2n=70

Smolen and Bickham (1994) Contributed by J. W. Bickham

ORDER RODENTIA

Dipodomys merriami (Kangaroo Rat)

2n=52

Mascarello et al. (1974) Contributed by A. D. Stock

2n=42

Graphodatsky (unpublished)

Sicista napaea (Birch Mouse)

187

188

EUARCHONTOGLIRES

Neotoma albigula (micropus) (Wood Rat/Pack Rat/Trade Rat)

2n=52

* Left=N. albigula

Mascarello et al. (1974)

Right=N. mictopus

Contributed by S. Pathak

Peromyscus eremicus (Cactus Mouse/White-Footed Mouse/Deer Mouse) G-bands

2n=48

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C-bands

2n=48

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Peromyscus maniculatus (White-Footed Mouse/Deer Mouse)

2n=48

Greenbaum et al. (1994)

Ideogram: Peromyscus maniculatus (White-Footed Mouse/Deer Mouse)

2n=48

Greenbaum et al. (1994)

189

190

EUARCHONTOGLIRES

Peromyscus boylii (Brush Mouse/White-Footed Mouse/Deer Mouse)

2n=48

Greenbaum (1994)

Oryzomys yunganus (Foothill Rice Rat)

2n=58

Volobouev and Aniskin (2000) Contributed by V. M. Aniskin

ORDER RODENTIA

191

Peromyscus truei (Piñon Mouse/White-Footed Mouse/Deer Mouse)

2n=48

Greenbaum et al. (1994)

Ideogram: Peromytcus truei (Piñon Mouse/White-Footed Mouse/Deer Mouse)

2n=48

Greenbaum et al. (1994)

192

EUARCHONTOGLIRES

Oryzomys nitidus (Elegant Rice Rat)

2n=80

Volobouev and Aniskin (2000) Contributed by V. M. Aniskin

Oryzomys megacephalus-Cytotype 2 (Rice Rat)

2n=54

Volobouev and Aniskin (2000) Contributed by V. M. Aniskin

ORDER RODENTIA

Oligoryzomys flavescens (Yellow Pygmy Rice Rat)

2n=66

Aniskin and Volobouev (1999)

Oligoryzomys microtis (Small-Eared Pygmy Rice Rat)

2n=64

Aniskin and Volobouev (1999)

193

194

EUARCHONTOGLIRES

Amphinectomys savamis (Water Rat) G-bands

2n=50

C-bands

2n=50

Malygin et al. (1994)

Malygin et al. (1994)

Contributed by V. M. Aniskin

Contributed by V. M. Aniskin

ORDER RODENTIA

Akodon simulator (Grey-Bellied Grass Mouse)

2n=36

Aniskin et al. (1996)

Akodon toba (Chako Grass Mouse)

2n=42

Aniskin et al. (1996)

195

196

EUARCHONTOGLIRES

Abrothrix (Akodon) sanborni (Sanborn’s Grass Mouse)

2n=52

Gallardo (1982)

Abrothrix (Akodon) longipilis (Long-Haired Grass Mouse)

2n=52

Gallardo (1982)

2n=52

Gallardo (1982)

Chroeomys (Akodon) olivaceus (Olive Grass Mouse)

ORDER RODENTIA

Eligmodontia puerulus (Highland Desert Mouse)

2n=50

Kelt et al. (1991) Contributed by M. H. Gallardo

Eligmodontia sp. (Highland Desert Mouse) Male

Female

2n=32

Kelt et al. (1991) Contributed by M. H. Gallardo

197

198

EUARCHONTOGLIRES

Maresomy (Auliscomys) boliviensis (Bolivian Big-Eared Mouse)

2n=22

Walker and Spotorno (1992) Contributed by A. E. Spotorno

Auliscomys sublimis (Andean Big-Eared Mouse)

2n=28

Walker and Spotorno (1992) Contributed by A. E. Spotorno

Loxodontomys (Auliscomys) micropus (Southern Big-Eared Mouse) Female

Male

2n=(34F) (32M)

Walker and Spotorno (1992) Contributed by A. E. Spotorno

ORDER RODENTIA

Euneomys chinchilloides petersoni (Patagonian Chinchilla Mouse)

2n=36

Reise and Gallardo (1990) Contributed by M. H. Gallardo

Euneomys mordax (Patagonian Chinchilla Mouse)

2n=42

Reise and Gallardo (1990) Contributed by M. H. Gallardo

199

200

EUARCHONTOGLIRES

Calomyscus mystax (Afghan Mouselike Hamster)

G-bands

2n=44

Radjabli, Sablina, and Graphodatsky (unpublished) Contriuted by A. Graphodatsky

C-bands

2n=44

Radjabli, Sablina, and Graphodatsky (unpublished) Contriuted by A. Graphodatsky

ORDER RODENTIA

Calomyscus urartensis (Urartsk Mouselike Hamster)

G-bands

2n=32

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C-bands

2n=32

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

201

202

EUARCHONTOGLIRES

Calomyscus bailwardi (Mouselike Hamster) G-bands

2n=44

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C-bands

2n=44

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Phodopus sungorus sungorus (Dzhungarian Hamster) G-bands

2n=28

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C-bands

2n=28

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

203

204

EUARCHONTOGLIRES

Phodopus sungorus campbelli (Campbell’s Hamster) G-bands

2n=28

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C-bands

2n=28

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Phodopus roborovskii (Small Desert Hamster) G-bands

2n=34

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C-bands

2n=34

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

205

206

EUARCHONTOGLIRES

Cricetus cricetus (Black-Bellied Hamster) G-bands

2n=22

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C-bands

2n=22

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Allocricetulus eversmanni (Eversmann’s Hamster) G - bands

2n=26

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C - bands

2n=26

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

207

208

EUARCHONTOGLIRES

Allocricetulus curtatus (Mongolian Hamster) G - bands

2n=20

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C - bands

2n=20

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Cricetulus migratorius (Gray Dwarf Hamster) G - bands

2n=22

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C - bands

2n=22

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

209

210

EUARCHONTOGLIRES

Cricetulus barabensis (Striped Dwarf Hamster)

G - bands

2n=20

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C - bands

2n=20

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Cricetulus pseudogriseus (Transbaikal Dwarf Hamster) G - bands

2n=24

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C - bands

2n=24

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

211

212

EUARCHONTOGLIRES

Cricetulus griseus (Chinese Hamster) G - bands

2n=22

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C - bands

2n=22

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Cricetulus obscurus (Gobi Hamster) G - bands

2n=20

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C - bands

2n=20

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

213

214

EUARCHONTOGLIRES

Cricetulus longicaudatus (Lesser Long-Tailed Hamster) G - bands

2n=24

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C - bands

2n=24

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Tscherskia ( Cricetulus ) triton (Greater Long-Tailed Hamster) G - bands

2n=28

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C - bands

2n=28

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

215

216

EUARCHONTOGLIRES

Mesocricetus brandti (Brandt’s Hamster) G - bands

2n=42

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C - bands

2n=42

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Mesocricetus brandti (Brandt's Hamster)

2n=44

Popescu and DiPaolo (1980) Contributed by N. Popescu

217

218

EUARCHONTOGLIRES

Mesocricetus auratus (Syrian Golden Hamster) G - bands

2n=44

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C - bands

2n=44

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Ideogram: Mesocricetus auratus (Syrian Golden Hamster)

2n=44

Popescu and DiPaolo (1972) Contributed by N. Popescu

219

220

EUARCHONTOGLIRES

Mesocricetus raddei (Ciscaucasian Hamster) G - bands

2n=44

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C - bands

2n=44

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Mesocricetus newtoni (Romanian Hamster) G - bands

2n=38

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

C - bands

2n=38

Radjabli, Sablina, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

221

222

EUARCHONTOGLIRES

Nannospalax (Spalax) ehrenbergi (Mediterranean Blind Mole-Rat)

2n=60

Ivanitskaya and Nevo (1998) Contributed by E. Nevo

Tatera nigricauda (Large Naked-Soled Gerbil)

2n=40

Qumsiyeh (1986)

ORDER RODENTIA

Tachyoryctes macrocephalus (Big-Headed Mole-Rat/African Mole-Rat)

G - bands

C - bands

2n=50

2n=50

Aniskin et al. (1997b)

Aniskin et al. (1997b)

223

224

EUARCHONTOGLIRES

Tachyoryctes splendens (East African Mole-Rat)

G - bands

C - bands

2n=48

2n=48

Aniskin et al. (1997b)

Aniskin et al. (1997b)

ORDER RODENTIA

Gerbillus rupicola (Rocky Gerbil/Northern Pygmy Gerbil)

2n=52

Granjon et al. (2002) Contributed by V. Volobouev

225

226

EUARCHONTOGLIRES

Taterillus petteri (Petter’s Gerbil/Small Naked-Soled Gerbil)

2n=19

Dobigny (2002)

Taterillus pygargus

(Senegal Gerbil/Small Naked-Soled Gerbil)

2n=22

Dobigny (2002)

ORDER RODENTIA

Taterillus arenarius (Senegal Gerbil/Small Naked-Soled Gerbil)

2n=35

Volobouev and Granjon (1996) Contributed by V. Volobouev

Taterillus tranieri (Tranier’s Gerbil/Small Naked-Soled Gerbil)

2n=15

Dobigny et al. (2003)

227

228

EUARCHONTOGLIRES

Taterillus congicus (Congo Gerbil/Small Naked-Soled Gerbil)

2n=54

Dobigny (2002)

ORDER RODENTIA

Taterillus sp. 1 (Small Naked-Soled Gerbil)

2n=23

Dobigny et al. (2002)

2n=23

Dobigny et al. (2002)

229

230

EUARCHONTOGLIRES

Taterillus sp. 2 (Small Naked-Soled Gerbil)

2n=37

Dobigny et al. (2002)

ORDER RODENTIA

Clethrionomys rufocanus (Gray Red-Backed Vole/Red-Backed Vole/ Bank Vole)

2n=56

Sokolov et al. (1990) Contributed by V. M. Aniskin

231

232

EUARCHONTOGLIRES

Clethrionomys sikotanensis (Sikotan Vole/Red-Backed Vole/Bank Vole)

2n=56

Sokolov et al. (1990) Contributed by V. M. Aniskin

ORDER RODENTIA

Clethrionomys rutilus (Northern Red-Backed Vole/Bank Vole)

2n=56

Sokolov et al. (1990) Contributed by V. M. Aniskin

233

234

EUARCHONTOGLIRES

Clethrionomys gapperi (Southern Red-Backed Mouse/Bank Vole)

2n=56

Modi (1987)

Clethrionomys californicus (Western Red-Backed Mouse/Bank Vole)

2n=56

Modi (1987)

ORDER RODENTIA

Clethrionomys glareolus (Red-Backed Vole/Bank Vole)

2n=56

Sokolov et al. (1990) Contributed by V. M. Aniskin

235

236

EUARCHONTOGLIRES

Clethrionomys centralis (Tien Shan Red-Backed Vole)

2n=56

Sokolov et al. (1990) Contributed by V. M. Aniskin

ORDER RODENTIA

Eothenomys regulus (Royal Vole/Pere David’s Vole/Pratt’s Vole)

2n=56

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=56

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

237

238

EUARCHONTOGLIRES

Alticola argentatus (Sliver High Mountain Vole)

2n=56

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=56

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Alticola macrotis (Large-Eared Vole)

2n=56

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=56

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

239

240

EUARCHONTOGLIRES

Phenacomys intermedius (Heather Vole)

2n=56

Modi (unpublished)

ORDER RODENTIA

Arvicola terrestris (European Water Vole/Bank Vole)

2n=36

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=36

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

241

242

EUARCHONTOGLIRES

Chionomys nivalis (European Snow Vole)

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Chionomys roberti (Robert’s Snow Vole)

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

243

244

EUARCHONTOGLIRES

Chionomys gud (Caucasian Snow Vole)

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Blanfordimys afghanus (Afghan Vole)

2n=58

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=58

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

245

246

EUARCHONTOGLIRES

Blanfordimys bucharicus (Bucharian Vole)

2n=48

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=48

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Microtus juldaschi carruthersi (Juniper Vole/Vole/Meadow Mouse)

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

247

248

EUARCHONTOGLIRES

Microtus subterraneus (European Pine Vole/Vole/Meadow Mouse)

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

249

Microtus daghestanicus (Daghestan Pine Vole/Vole/Meadow Mouse)

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

250

EUARCHONTOGLIRES

Microtus majori (Major’s Pine Vole/Vole/Meadow Mouse)

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Microtus nasarovi (Nasarov’s Vole/Vole/Meadow Mouse)

2n=42

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=42

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

251

252

EUARCHONTOGLIRES

Microtus schelkovnikovi (Schelkovnikov’s Pine Vole/Vole/Meadow Mouse)

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Microtus pinetorum (Campagnol/Sylvestre Pine Vole/Woodland Vole/Vole/Meadow Mouse)

2n=62

Modi (1987)

Microtus ochrogaster (Prairie Vole/Vole/Meadow Mouse)

2n=54

Modi (1987)

253

254

EUARCHONTOGLIRES

Microtus richardsoni (Water Vole/Vole/Meadow Mouse)

2n=56

Modi (1987)

Microtus longicaudus (Coronation Island Vole/Long-Tailed Vole/ Vole/Meadow Mouse)

2n=64

Modi (1987)

ORDER RODENTIA

Microtus socialis (Social Vole/Vole/Meadow Mouse)

2n=62

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=62

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

255

256

EUARCHONTOGLIRES

Microtus socialis schiddlovskii (Schiddlovskii’s Social Vole/Vole/Meadow Mouse)

2n=60

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

Microtus guentheri arm (Günther’s Vole/Vole/Meadow Mouse)

2n=60

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Microtus guentheri philistinus (Günther’s Vole/Vole/Meadow Mouse)

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

257

258

EUARCHONTOGLIRES

Microtus arvalis arvalis (Common Vole/Vole/Meadow Mouse)

2n=46

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=46

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Microtus rossiaemeridionalis (Southern Vole/Vole/Meadow Mouse)

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

259

260

EUARCHONTOGLIRES

Microtus transcaspicus (Transcaspian Vole/Vole/Meadow Mouse)

2n=52

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=52

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Microtus kirgisorum (Tien Shan Vole/Vole/Meadow Mouse)

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

261

262

EUARCHONTOGLIRES

Microtus oeconomus (Tundra Vole/Vole/Meadow Mouse)

2n=30

Modi (1987)

Microtus montebelli (Japanese Grass Vole/Vole/Meadow Mouse)

2n=30

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Microtus fortis (Reed Vole)

2n=52

Kovalskaya et al. (1991) Contributed by V. M. Aniskin

263

264

EUARCHONTOGLIRES

Microtus mongolicus (Mongolian Vole/Vole/Meadow Mouse)

2n=50

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

Microtus hyperboreus (North Siberian Vole/Vole/Meadow Mouse)

2n=50

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Microtus sachalinensis (Sakhalin Vole/Vole/Meadow Mouse)

2n=50

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=50

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

265

266

EUARCHONTOGLIRES

Microtus maximowiczii (Maximowicz’s Vole/Vole/Meadow Mouse)

2n=44

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

Microtus mujanensis (Muisk Vole/Vole/Meadow Mouse)

2n=38

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Microtus evoronensis (Evoronsk Vole/Vole/Meadow Mouse)

2n=40

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=40

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

267

268

EUARCHONTOGLIRES

Microtus gregalis (Narrow-Headed Vole/Vole/Meadow Mouse)

2n=36

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=36

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Microtus agrestis (Field Vole/Vole/Meadow Mouse)

2n=50

Microtus pennsylvanicus (Campagnol des Champs/Meteoro de Prado/ Vole/Meadow Mouse)

Modi (1987)

2n=46

Modi (1987)

269

270

EUARCHONTOGLIRES

Microtus breweri (Beach Vole/Vole/Meadow Mouse)

2n=46

Modi (1987)

Microtus townsendii (Townsend’s Vole/Vole/Meadow Mouse)

2n=50

Modi (1987)

ORDER RODENTIA

Microtus oregoni (Creeping Vole/Vole/Meadow Mouse)

2n=17–18

Modi (1987)

Microtus californicus (Amargosa Vole/California Vole/Vole/ Meadow Mouse)

2n=53

Modi (1987)

271

272

EUARCHONTOGLIRES

Microtus mexicanus (Mexican Vole/Vole/Meadow Mouse)

2n=48

Modi (1987)

Lagurus lagurus (Steppe Lemming)

2n=56

Modi (1987)

ORDER RODENTIA

Microtus sp. nova from lran (Vole/Meadow Mouse)

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=54

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

273

274

EUARCHONTOGLIRES

Neofiber alleni (Round-Tailed Muskrat/Florida Water Rat)

2n=52

Modi (1987)

Dicrostonyx torquatus (Collared Lemming/Varying Lemming)

2n=30

Modi (1987)

ORDER RODENTIA

Lemmus sibiricus (True Lemming)

2n=50

Modi (1987)

Synaptomys cooperi (Bog Lemming)

2n=50

Modi (1987)

275

276

EUARCHONTOGLIRES

Prometheomys schaposchnikowi (Long-Clawed Mole-Vole)

2n=56

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Ellobius talpinus (Northern Mole-Vole/Mole-Lemming)

2n=50

Perelman and Graphodatsky (unpublished) Contributed by A. Graphodatsky

277

278

EUARCHONTOGLIRES

Ellobius fuscocapillus (Southern Mole-Vole/Mole-Lemming)

2n=36

Perelman and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=36

Perelman and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Ellobius lutescens (Transcaucasian Mole-Vole/Mole-Lemming)

2n=17

Perelman Graphodatsky, Baklushinskaya, and Lyapunova (unpublished) Contributed by A. Graphodatsky

Apodemus peninsulae (Korean Wood Mouse/Field Mouse/ Old World Wood Mouse)

2n=48(+B’S)

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

279

280

EUARCHONTOGLIRES

Apodemus sylvaticus (Long-Tailed Field Mouse/Field Mouse/Old World Wood Mouse)

2n=48

Stanyon et al. (2004)

ORDER RODENTIA

Rattus rattus (House Rat/Black Rat/Roof Rat)

2n=38

Sablina and Graphodatsky (unpublished) Contributed by A. Graphodatsky

281

282

EUARCHONTOGLIRES

Rattus norvegicus (Norway Rat/Brown Rat)

2n=42

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Rattus flavipectus (Buff-Bellied Rat)

2n=46

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

283

284

EUARCHONTOGLIRES

Phloeomys cumingi (Slender-Tailed Cloud Rat)

2n=44

Rickart and Heany (2002)

Crunomys suncoides (Philippine Rat/Sulawesian Shrew Rat)

Contributed by Rickart

2n=36

Rickart and Heany (2002) Contributed by Rickart

ORDER RODENTIA

Archboldomys luzonensis (Mt Isaroq Shrew Rat)

2n=26

Rickart and Heany (2002) Contributed by Rickart

Celaenomys silaceus (Luzor Striped Rat)

2n=44

Rickart and Heany (2002) Contributed by Rickart

285

286

EUARCHONTOGLIRES

Nesokia indica (Short-Tailed Bandicoot Rat/Pest Rat)

2n=42

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Leopoldamys edwardsi (Long-Tailed Giant Rat)

2n=44

Sablina, Radjabli, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

287

288

EUARCHONTOGLIRES

Mastomys natalensis (Multimammate Rat)

2n=32

2n=32

Volobouev et al. (2002a)

Volobouev et al. (2002a)

ORDER RODENTIA

Mastomys erythroleucus-Cytotype 3 (Guinea Multimammate Rat)

2n=38

2n=38

Volobouev et al. (2002a)

Volobouev et al. (2002a)

289

290

EUARCHONTOGLIRES

Mastomys awashensis (Multimammate Rat)

2n=32

2n=32

Volobouev et al. (2002a)

Volobouev et al. (2002a)

ORDER RODENTIA

Mastomys huberti (Multimammate Rat)

2n=32

2n=32

Volobouev et al. (2002a)

Volobouev et al. (2002a)

291

292

EUARCHONTOGLIRES

Mastomys kollmanspergeri (Kollmansperger’s Multimammate Rat)

2n=38

2n=38

Volobouev et al. (2002a)

Volobouev et al. (2002a)

ORDER RODENTIA

Myomys daltoni (Dalton’s Mouse)

2n=36

2n=36

Volobouev et al. (2002a)

Volobouev et al. (2002a)

293

294

EUARCHONTOGLIRES

Stenocephalemys albocaudata (Ethiopian Narrow-Headed Rat)

2n=54

Lavrenchenko et al. (2001) Contributed by V. M. Aniskin

ORDER RODENTIA

Stenocephalemys griseicauda (Gray-Tailed Narrow-Headed Rat)

2n=54

Lavrenchenko et al. (2001) Contributed by V. M. Aniskin

295

296

EUARCHONTOGLIRES

Stenocephalemys albipes (Ethiopian Narrow-Headed Rat)

2n=46

Lavrenchenko et al. (2001) Contributed by V. M. Aniskin

ORDER RODENTIA

297

Lophuromys melanonyx (Black-Clawed Brush-Furred Rat/Brush-Furred Mouse)

2n=60

Aniskin et al. (1997a)

298

EUARCHONTOGLIRES

Lophuromys brevicaudus (Short-Tailed Brush-Furred Rat/Brush-Furred Mouse)

2n=68

Aniskin et al. (1997a)

ORDER RODENTIA

Lophuromys chrysopus (Brush-Furred Rat/Brush-Furred Mouse)

2n=54

Aniskin et al. (1997a)

299

300

EUARCHONTOGLIRES

Dasymys incomtus (African Marsh Rat/Shaggy Swamp Rat)

2n=36

2n=36

Volobouev et al. (2000)

Volobouev et al. (2000)

ORDER RODENTIA

301

Dasymys rufulus (West African Shaggy Rat/Shaggy Swamp Rat)

2n=36+B’s

2n=36+B’s

Volobouev et al. (2000)

Volobouev et al. (2000)

302

EUARCHONTOGLIRES

Arvicanthis niloticus-Cytotype 1a (African Grass Rat/Unstriped Grass Rat/Kusu Rat)

2n=62

2n=62

Volobouev et al. (2002b)

Volobouev et al. (2002b)

ORDER RODENTIA

Arvicanthis niloticus-Cytotype 1b (African Grass Rat/Unstriped Grass Rat/Kusu Rat)

2n=62

2n=62

Volobouev et al. (2002b)

Volobouev et al. (2002b)

303

304

EUARCHONTOGLIRES

Arvicanthis ansorgei (African Grass Rat/Unstriped Grass Rat/Kusu Rat)

2n=62

2n=62

Volobouev et al. (2002b)

Volobouev et al. (2002b)

ORDER RODENTIA

Arvicanthis rufinus (African Grass Rat/Unstriped Grass Rat/Kusu Rat)

2n=62

2n=62

Volobouev et al. (2002b)

Volobouev et al. (2002b)

305

306

EUARCHONTOGLIRES

Arvicanthis sp. (African Grass Rat/Unstriped Grass Rat/Kusu Rat)

2n=58

2n=58

Volobouev et al. (2002b)

Volobouev et al. (2002b)

ORDER RODENTIA

Mus musculus (House Mouse)

2n=40

Stanyon (unpublished)

307

308

EUARCHONTOGLIRES

Ideogram: Mus musculus (House Mouse)

2n=40

Adler (1994)

ORDER RODENTIA

Mus macedonicus (Macedonian Mouse)

2n=40

Ivanitskaya et al. (1996) Contributed by E. Nevo

Mus mahomet (Mahomet Mouse)

2n=36

Aniskin et al. (1998)

309

310

EUARCHONTOGLIRES

Pedetes capensis (Springhare/Springhaas)

2n=38

Biltueva and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Pectinator spekei (Speke’s Pectinator)

2n=40

George (1979)

2n=36

George (1979)

Massoutiera mzabi (Massoutiera)

311

312

EUARCHONTOGLIRES

Ctenodactylus gundi (Atlas Gundi)

2n=40

George (1979)

2n=40

George (1979)

Ideogram: Ctenodactylus gundi (Atlas Gundi)

ORDER RODENTIA

Ctenodactylus vali (Sahara Gundi)

2n=40

George (1979)

313

314

EUARCHONTOGLIRES

Muscardinus avellanarius (Common Dormouse/Hazel Mouse)

2n=46

Graphodatsky (unpublished)

ORDER RODENTIA

Myoxus glis (Fat Dormouse/Edible Dormouse)

2n=62

Graphodatsky (unpublished)

315

316

EUARCHONTOGLIRES

Dryomys nitedula (Forest Dormouse)

2n=48

Graphodatsky (unpublished)

ORDER RODENTIA

Eliomys quercinus (Garden Dormouse)

2n=48

Graphodatsky (unpublished)

317

318

EUARCHONTOGLIRES

Myomimus personatus (Mouselike Dormouse)

2n=44

Graphodatsky (unpublished)

ORDER RODENTIA

Georychus capensis (Cape Mole-Rat)

2n=54

Nevo et al. (1986)

2n=54

Nevo et al. (1986)

Bathyergus janetta (Dune Mole-Rat)

319

320

EUARCHONTOGLIRES

Cryptomys mechowi (Giant Mole-Rat/Common Mole-Rat)

2n=40

Macholan et al. (1993)

ORDER RODENTIA

Cryptomys hottentotus damarensis (Common Mole-Rat)

2n=78

Nevo et al. (1986)

*From Kalahari desert -high number subpecies

321

322

EUARCHONTOGLIRES

Lagostomus maximus (Plains Viscacha)

2n=56

Graphodatskaya and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Dinomys branickii (Pacarana)

2n=58

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

323

324

EUARCHONTOGLIRES

Cavia porcellus (cobaya) (Guinea Pig/Cavies)

2n=64

Graphodatsky and Sablina (unpublished) Contributed by A. Graphodatsky

ORDER RODENTIA

Hydrochaeris hydrochaeris (Capybara)

2n=66

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Ctenomys rionegrensis (Tuco-Tuco)

2n=50

L. Garcia et al. (2002) Contributed by M. Ponsa

325

326

EUARCHONTOGLIRES

Ctenomys talarum (Tuco-Tuco)

2n=48

Ctenomys maulinus brunneus Contributed by M. Ponsa (Tuco-Tuco)

L. Garcia et al. (2002)

2n=26

Gallardo (1979)

Ctenomys robustus (Tuco-Tuco)

2n=26

Gallardo (1979)

ORDER RODENTIA

Ctenomys maulinus maulinus (Tuco-Tuco)

2n=26

Gallardo (1979)

Ctenomys magellanicus magellanicus (Tuco-Tuco)

2n=34

Gallardo (1979)

Ctenomys opimus (Tuco-Tuco)

2n=26

Gallardo (1979)

327

328

EUARCHONTOGLIRES

Ctenomys fulvus (Tuco-Tuco)

2n=26

Gallardo (1979)

Octodon lunatus (Degus)

2n=78

Spotorno et al. (1995)

ORDER RODENTIA

Octodon degus (Degus)

2n=58

Spotorno et al. (1995)

329

330

EUARCHONTOGLIRES

Tympanoctomys barrerae (Viscacha Rat)

2n=102

Gallardo et al. (2004)

ORDER RODENTIA

Tympanoctomys barrerae (Viscacha Rat)

2n=102

Gallardo et al. (2004)

C-bands This is an all-biarmed karyotype. The Y chromosome is the only acrocentric of the whole complement. Note the centromeric position of heterochromatin blocks in all chromosomes regardless of shape. Only one pair having the secondary constriction (SC) is observed.

331

332

EUARCHONTOGLIRES

Spalacopus cyanus (Coruro)

2n=58

Spotorno et al. (1995)

ORDER RODENTIA

Aconaemys fuscus (Rock Rat)

2n=56

Gallardo and Mondaca (2002) Contributed by Gallardo

Aconaemys sagei (Rock Rat)

2n=54

Gallardo and Mondaca (2002) Contributed by Gallardo

Aconaemys porteri (Rock Rat)

2n=58

Gallardo and Mondaca (2002) Contributed by Gallardo

333

334

EUARCHONTOGLIRES

Abrocoma bennetti (Chinchilla Rat/Chinchillone)

2n=64

Spotorno et al. (1995)

ORDER RODENTIA

Proechimys brevicauda-Cytotype 1 (Huallaga Spiny Rat/Casiragua)

2n=28

Aniskin (1994b)

2n=28

Aniskin (1994b)

335

336

EUARCHONTOGLIRES

Proechimys steerei (Sterre’s Spiny Rat/Casiragua)

2n=24

Aniskin (1994b)

2n=24

Aniskin (1994b)

ORDER RODENTIA

Proechimys simonsi (Simon’s Spiny Rat/Casiragua)

2n=32

Aniskin (1994b)

2n=32

Aniskin (1994b)

337

338

EUARCHONTOGLIRES

Proechimys sp. 1 (Terrestrial Spiny Rat/Casiragua)

2n=30

Aniskin (1994b)

Proechimys sp. 2 (Terrestrial Spiny Rat/Casiragua)

2n=34

Aniskin (1994b)

ORDER RODENTIA

Lonchotrix emiliae (Tuft-Tailed Spiny Tree Rat)

2n=62

Aniskin (1994b)

Isothrix sinnamariensis (French Guiana Brush-Tailed Rat/Toros)

2n=28

Vié et al. (1996) Contributed by V. Volobouev

339

340

EUARCHONTOGLIRES

Dactylomys dactylinus (Amazon Bamboo Rat/Coro-Coro)

2n=96

Aniskin (1993)

ORDER RODENTIA

Myocastor coypus (Nutria/Coypu)

2n=42

Biltueva and Graphodatsky (unpublished) Contributed by A. Graphodatsky

341

342

EUARCHONTOGLIRES

Order Lagomorpha The order Lagomorpha comprises two families, Leporidae (hares and rabbits, with 11 genera and approximately 56 species; Angerman et al., 1990) and Ochotonidae (pikas, 1 genus and some 26 species; Angerman et al., 1990; Nowak, 1999; Yu et al., 2000). Lagomorphs sensu latu have a natural distribution that includes the Holarctic, Ethiopian (excluding Madagascar), Northern Neotropica, and northern and western Oriental regions (DeBlase and Martin, 1981). Although there is now substantial evidence to suggest a sister relationship with rodents (Glires concept; Gregory, 1910; Simpson, 1945), lagomorphs have, in the past, also been grouped with elephant shrews in Anagalida (Lagomorpha + Rodentia + Macroscelidea; McKenna and Bell, 1997; Shoshani and McKenna, 1998). However, concatenated sequences from mitochrodrial and nuclear genes provide overwhelming support for elephant shrews to group within Afrotheria, a clade of probable African origin that also contains hyraxes, sirenians, elephants, aardvarks, tenrecs, and golden moles (Springer et al., 1997; M. J. Stanhope et al., 1998; Amrine-Madsen et al., 2003; Murphy et al., 2001a,b).

Family Leporidae In terms of generic distribution, the almost ubiquitous occurrence of the specious Lepus contrasts markedly with most of the remaining taxa which are monotypic (Brachylagus, Pentalagus, Caprolagus, Bunolagus, Poelagus, Romerolagus, and Oryctolagus), several of which show narrow endemism prompting their recognition as endangered or vulnerable in the International Union for Conservation of Nature (IUCN) categories of threat (Chapman and Flux, 1990). The only exceptions to monotypic status are Nesolagus (2 species; Surridge et al., 1999; Can et al., 2001), Sylvilagus (16 species; Angerman et al., 1990; Chapman et al., 1992; Frey et al., 1997), and Pronolagus (4 species; Whiteford, 1995; Matthee and Robinson, 1996). Obtaining a robust phylogeny for Leporidae has proved elusive with morphology (Corbet, 1983, and references therein), cytogenetics (Robinson et al., 2002, and references therein), and mitochondrial sequences (Halanych and Robinson, 1999) plagued by convergence (morphology), autapomorphic characters (cytogenetics), or excessive homoplasy (sequences). Most recently, analysis of a supermatrix constructed for 27 taxa representing all 11 leporid genera (5483 characters from five nuclear and two mtDNA gene fragments) grouped the extant leporids into two evolutionary clades (Matthee et al., 2004). One clade comprised the African Poelagus and Pronolagus as sister taxa, with the Nesolagus basal to these; the second clade included Romerolagus, Lepus, Sylvilagus, Brachylagus, Pentalagus, Bunolagus, Caprolagus, and Oryctolagus. Within this assemblage, the New World cottontail genera Brachylagus and Sylvilagus are sister taxa. There was unambiguous support for the monophyly of a derived group comprising Pentalagus, Bunolagus, Caprolagus, and Oryctolagus, but the precise evolutionary relationships among them was not resolved. Chromosome numbers within Leporidae range from 2n = 38 to 2n = 52 (Holden, 1968; Robinson et al., 1983). However, the evolutionary groupings suggested by the Matthee et al. (2004) supermatrix analysis were not reflected in the earlier G-banding studies (Robinson et al., 1984, and references therein) or by the more recent fluorescence in situ hybridization (FISH) data (Robinson et al., 2002). Cross-species chromosome painting using flow-sorted painting probes developed from the Old World rabbit (Korstanje et al., 1999) showed that at least 18 fusions and 6 fissions are required to derive the

ORDER LAGOMORPHA karyotypes of 7 of the 11 extant genera from the presumed ancestral state (2n = 48). Although one rearrangement (a centric fusion) is shared by Oryctolagus and Brachylagus, there is no support for a close evolutionary affinity of these taxa from either morphology or DNA sequences, and the most likely explanation is that the arrangement arose independently in both lineages. In contrast to the intergeneric comparisons, however, a comparative cytogenetic study of a subset of Sylvilagus species identified chromosomal states that were phylogenetically informative, suggesting evolutionary associations within the cottontails that are consistent with those predicted by mtDNA sequence data (Halanych and Robinson, 1997). It has been suggested that leporid chromosomal evolution probably arose in response to some form of spatial isolation, and the failure of the chromosomal data to provide markers that track descent from common ancestry reflects the group’s rapid radiation. In addition to the evolutionary studies referred to above, complete homology maps between Oryctolagus cuniculus and humans have been established by reciprocal chromosome painting. Using 22 rabbit-specific (O. cuniculus) chromosome-painting probes, Korstanje and colleagues (1999) identified 42 homologous segments when these were hybridized to humans, with 38 segments evident in the reciprocal direction. The delineation of conserved syntenic segments between rabbit and human chromosomes facilitated the regional localization of loci on the gene-poor rabbit chromosomes since these could often be predicted by the comparative maps. For example, Zijlstra et al. (2002) and Hayes et al. (2002) were able to assign in excess of 30 markers to the rabbit map and to correct and refine some of the earlier painting data. Moreover, the localization of several microsatellite loci (Zijlstra et al., 2002) has made it possible to assign linkage groups to individual rabbit chromosomes, thus contributing significantly to the development of an integrated cytogenetic and genetic map for this species (see the INRA Rabbitmap database at http://locus.jouy.inra.fr/cgi-bin/lgbc/mapping/common/intro2.pl? BASE=rabbit).

Family Ochotonidae Pikas occur only in the northern hemisphere and are partitioned into the nonburrowing tallus species and burrowing species that occur in steppe, shrub, and forest habitat (see Smith et al., 1990, and references therein). The most recent comprehensive phylogenetic investigation of the ochotonids was based on 19 species and 23 forms (taxa of questionable status) using mtDNA cytochrome b and ND4 gene sequences (Yu et al., 2000). Their data show three major clades represented by the shrub-steppe species, a northern group and a mountain group. A biogeographic interpretation of their results suggests that vicariance, promoted by uplifting of the Tibet (Qinghai-Xizang) Plateau, had a major influence on cladogenesis. In contrast, the nearctic Ochotona princeps probably arose through dispersion during the Pleistocene opening of the Bering Straight. Comparative cytogenetic data on Ochotonidae are available for Ochotona pusilla (2n = 68), O. rutila (2n = 62), O. rujescens (2n = 50), O. daurica (2n = 50), O. alpina (2n = 42), and O. pallasi (2n = 38; Ivanitskaya, 1991). C-banding patterns have also been published for O. hyperborea (2n = 40; Ivanitskaya, 1991). In addition to these taxa, G- and C-banded chromosomes of the nearctic O. princeps (2n = 68; Stock, 1976) and the unbanded chromosomes of O. collaris (2n = 68; Hsu and Benirschke, 1971) have been published. A comparative analysis of pika G-banded chromosomes by Ivanitskaya (1991) shows that O. alpina and O. pallasi and O. pusilla and O. princeps are, respectively, karyotypically most similar, tracking correspondence in their diploid

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EUARCHONTOGLIRES numbers. Interestingly, although far from having the taxon representation of the molecular phylogenetic study (Yu et al., 2000), the chromosomes suggest the presence of two evolutionary clades (subgenera) within Ochotona. These are O. pusilla, O. princeps, O. rutila, O. rujescenc, and O. daurica, which are referable to Ochotona (and which may also include Ochotona collaris, O. macrotis, and O. curzoinae), and Ochotona alpina, O. pallasi, and O. hyperborean, which are grouped within the subgenus Pika. However, agreement on the composition of the clades suggested by the chromosomal data on one hand and the sequences on the other is weak. In large part this may be due to so few species in common to the studies, and a rigorous assessment of correspondence will be dependent on more equitable species representation. Comparative G-banding between O. princeps and O. cuniculus by Stock (1975) showed that only a small portion of the Ochotona genome is directly homologous on banding pattern to any of the Oryctolagus chromosomes, suggesting that intrachromosomal rearrangements have figured prominently in the evolution of the two major lagomorph lineages. Most recently Graphodatsky and Yang (Personal Communication) identified 40 conserved autosomal segments between human and O. hyperborea using human chromosome-painting probes. A comparison of the human–rabbit (Korstanje et al., 1999) and human–ochotonid (Grapohodatsky and Yang (Personal Communication)) chromosome maps shows that most syntenic associations identified in O. cuniculus are also present in O. hyperborea (sometimes with a new fusion partner), suggesting that Robertsonian changes have driven the differences in diploid numbers and an examination of G-banding pattern homology between lower numbered ochotonids and the leporids may detect extensive regions of homology among them. Terence J. Robinson

ORDER LAGOMORPHA

Ochotona princeps (American Pika/Mouse Hare/Conie)

2n=68

Stock (1976)

2n=68

Stock (1976)

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EUARCHONTOGLIRES

Ochotona hyperborea (Northern Pika/Mouse Hare/Conie)

2n=40

Perelman and Graphodatsky (unpublished) Contributed by A. Graphodatsky

Pronolagus rupestris (Red Rabbit)

2n=42

Robinson et al. (2002)

ORDER LAGOMORPHA

Bunolagus monticularis (Bushman Rabbit)

2n=44

Robinson et al. (2002)

347

348

EUARCHONTOGLIRES

Romerolagus diazi (Volcano Rabbit)

2n=48

Robinson et al. (2002)

ORDER LAGOMORPHA

Brachylagus idahoensis (Pygmy Rabbit)

2n=44

Robinson et al. (2002)

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EUARCHONTOGLIRES

Sylvilagus aquaticus (Swamp Rabbit/Cottontail)

2n=38

Robinson et al. (1983)

Sylvilagus palustris (Marsh Rabbit/Cottontail)

2n=38

Robinson et al. (1983)

ORDER LAGOMORPHA

Sylvilagus transitionalis (New England Cottontail)

2n=46

Robinson et al. (1983)

Sylvilagus floridanus (Eastern Cottontail)

2n=42

Robinson et al. (1983)

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EUARCHONTOGLIRES

Sylvivagus nuttallii (Mountain Cottontail)

2n=42

Robinson et al. (1984)

2n=42

Robinson et al. (1984)

Sylvivagus audubonii (Desert Cottontail)

ORDER LAGOMORPHA

Lepus californicus (Black-Tailed Jack Rabbit/Hare)

2n=48

Robinson et al. (2002)

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EUARCHONTOGLIRES

Lepus saxatilis (Jack Rabbit/Hare)

2n=48

Robinson (1980)

Oryctolagus cuniculus (Domestic Rabbit/Old World Rabbit)

2n=44

Robinson et al. (2002)

ORDER LAGOMORPHA

Ideogram: Oryctolagus cuniculus (Domestic Rabbit/Old World Rabbit)

2n=44

Hayes et al. (2002)

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LAURASIATHERIA Order Eulipotyphla The monophyletic order Eulipotyphla comprises approximately 450 living species classified in four families: Solenodontidae (the solenodons), Talpidae (the moles and desmans), Erinaceidae (the hedgehogs and gymnures), and Soricidae (the shrews). Eulipotyphlans have traditionally been classified in an inclusive order “Insectivora” with a number of other mammalian orders, including tenrecs and golden moles (order Afrosoricida), elephant shrews (order Macroscelida), and tree shrews (order Scandentia). Though phenetically similar to eulipotyphlan families in certain traits (mostly reflecting shared plesiomorphies), it is now known that the evolutionary origins of these various orders are diverse and that among placental mammals they are not closely allied to eulipotyphlans (M. J. Stanhope et al., 1998; Murphy et al., 2001b; Clark et al., 2001). Cytogenetic aspects of eulipotyphlan families were discussed by Borgaonkar (1969), Gropp (1969), and Fredga (1970) and are reviewed in detail on a useful website maintained by the Institute of Cytology and Genetics, Novosibirsk (www.bionet.nsc.ru/chromosomes). In general, chromosomes of most eulipotyphlans have received little or moderate scientific attention. However, the cytogenetics of one particular taxon (the Sorex araneus species complex) have been studied in considerable detail because of two interesting evolutionary properties: a sex determination system (XX/XY1Y2) unique among eulipotyphlans (Fredga, 1970) and extensive chromosomal polymorphism among various karyotypic “races” (Hausser, 1991). Cytogenetic aspects of each eulipotyphlan family are briefly reviewed here.

Family Solenodontidae 2n = 34 (XX/XY?): Solenodons This endemic West Indian family comprises two living species, both classified in monotypic genera—the Cuban solenodon (Atopogale cubanus), endemic to Cuba, and the Hispaniolan solenodon (Solenodon paradoxus), endemic to Hispaniola (Roca et al., 2004). Both are highly endangered species. Solenodons represent the sister group to all other eulipotyphlans (Roca et al., 2004) and are anatomically the most primitive living members of the order in many ways (Asher, 2001).

Atlas of Mammalian Chromosomes, Edited by Stephen J. O’Brien, Joan C. Menninger, William G. Nash Copyright © 2006 John Wiley & Sons, Inc.

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LAURASIATHERIA Only a single individual S. paradoxus (female) has been karyotyped (Borgaonkar, 1969), revealing a relatively low diploid number (2n = 34), with fundamental number (FN) 64. The sex determination system is assumed to be XX/XY, as in most members of the order, but this requires verification. Given their deep molecular divergence (Roca et al., 2004), cytogenetic comparisons between the two extant solenodon genera would be of particular biological interest.

Family Talpidae 2n = 32–42 (XX/XY): Moles, shrew-moles, and desmans Talpids (the moles, shrew-moles, and desmans) are a group of fossorial and semiaquatic species largely restricted to temperate regions in North America and Eurasia. Talpids have traditionally been arranged as the sister group to the shrews (Soricidae), and the two families have been united in the order Soricomorpha (McKenna and Bell, 1997; Hutterer, 2005). More recent analyses, however, suggest that shrews and hedgehogs are actually more closely related, to the exclusion of talpids (Roca et al., 2004). Within Talpidae, current taxonomy recognizes approximately 40 living species in 17 genera and 3 subfamilies—Talpinae, Scalopinae, and Uropsilinae (Hutterer, 2005). Relevant reviews of issues in talpid cytogenetics include papers by Gropp (1969), Reumer and Meylan (1986), Harada et al. (2001), and Kawada et al. (2002). Representatives of at least 11 talpid genera have been karyotyped. As a family, talpids are moderately variable in both diploid number and the structural organization of the karyotype. Interestingly, the two species of desmans (semiaquatic talpids endemic to Europe—subfamily Talpinae, tribe Desmanini) have the lowest and highest diploid complements recorded in the family, respectively (2n = 32 in Desmana moschata; 2n = 42 in Galemys pyrenaicus). Most other talpids (i.e., moles and shrew-moles) have a diploid number of 2n = 34–38. Of the three talpid subfamilies, only Uropsilinae remains unstudied cytogenetically. The sex determination system is XX/XY.

Family Erinaceidae 2n = 48 (XX/XY): Hedgehogs and gymnures Erinaceidae can be divided into two distinctive subfamilies—Erinaceinae, comprising the hedgehogs (15–20 species in 4 genera), and Galericinae, the gymnures (approximately 7 species in 3–5 genera). Hedgehogs are distributed in relatively dry and open habitats throughout Africa and Eurasia, while gymnures are endemic to the tropical and subtropical forests of east and southeast Asia. The chromosomes of many hedgehog species have been studied, but gymnures remain cytogenetically unknown, suggesting a priority for future research. The diploid number appears to be 2n = 48 in all hedgehogs studied to date, although there is an early (incorrect?) report of 2n = 44 in Korean Erinaceus amurensis (see Borgaonkar, 1969). Despite this consistency in number, comparisons between species reveal considerable variation in karyotypic structure (e.g., Gropp, 1969). Mandahl (1979) discussed the position of nucleolar organizer regions (NORs). The sex determination system is XX/XY.

Family Soricidae 2n = 20—68 (XX/XY, XX/XY1Y2): Shrews The shrews (family Soricidae) constitute one of the largest, most widespread, and most successful mammalian families, with almost 400 species in approxi-

ORDER EULIPOTYPHLA mately 30 genera. Shrews are distributed throughout Eurasia, Africa, North America, and northern South America across a wide range of habitats, including arctic, temperate, and tropical biomes. The chromosomes of shrews have received more attention than in most other groups of mammals, and an extensive cytogenetic literature is available, especially with regard to the genera Sorex, Crocidura, Suncus, and Blarina. Diploid numbers vary widely among shrews, from 2n = 20 (some Sorex) to 2n = 68 (Notiosorex cf. crawfordi and Crocidura yankariensis; see Baker and Hsu, 1970; Schlitter et al., 1999), as do fundamental numbers. Recent cytogenetic studies of shrews have centered largely on studies of speciation and species-level boundaries, chromosomal polymorphism, and comparative genome mapping (e.g., Lukacova et al., 1996; Elrod et al., 1996; Larkin et al., 2000; Biltueva et al., 2001; Matsubara et al., 2001). As noted above, considerable historical scientific attention has focused on sex determination and chromosomal polymorphism in the Sorex araneus species-group (reviewed by Hausser, 1991). Kristofer M. Helgen Don Wilson

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LAURASIATHERIA

Podogymnura truei (Philippine Gymnure/Philippine Wood-Shrew)

2n=40

Rickart (2003)

Erinaceus europaeus (Western Eurasian Hedgehog)

2n=48

Radjabli and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER EULIPOTYPHLA

Erinaceus concolor (Eastern Eurasian Hedgehog)

2n=48

Sokolov et al. (1991) Contributed by V. M. Aniskin

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362

LAURASIATHERIA

Erinaceus amurensis (Eurasian Hedgehog/Amur Hedgehog)

2n=48

Radjabli and Graphodatsky (unpublished) Contributed by A. Graphodatsky

2n=48

Radjabli and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER EULIPOTYPHLA

Paraechinus hypomelas (Brandt’s Hedgehog/Desert Hedgehog)

2n=48

Radjabli and Graphodatsky (unpublished) Contributed by A. Graphodatsky

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364

LAURASIATHERIA

Hemiechinus auritus (Long-Eared Desert Hedgehog)

2n=48

Radjabli and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER EULIPOTYPHLA

Mesechinus (Hemiechinus) dauricus (Daurian Hedgehog)

2n=48

Radjabli and Graphodatsky (unpublished) Contributed by A. Graphodatsky

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LAURASIATHERIA

Solenodon paradoxus (Hispaniolan Solenodon)

2n=34

Benirschke (1969)

Sorex minutus (Eurasian Pygmy Shrew/Long-Tailed Shrew)

2n=42

Perelman, Biltueva, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER EULIPOTYPHLA

Sorex caecutiens (Laxmann’s Shrew/Long-Tailed Shrew)

2n=42

Perelman, Biltueva, and Graphodatsky (unpublished)

Sorex raddei (Radde’s Shrew/Long-Tailed Shrew)

Contributed by A. Graphodatsky

2n=36

Perelman, Biltueva, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

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368

LAURASIATHERIA

Sorex araneus (Eurasian Shrew/Long-Tailed Shrew)

2n=21

Perelman, Biltueva, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

Ideogram: Sorex araneus (Eurasian Shrew/Long-Tailed Shrew)

2n=21

Serov et al. (1998)* * “Reprinted with permission from ILAR Journal, 39: 195–202 (1998), Institute for Laboratory Animal Research, The Keck Center of the National Academies, 500 Fifth Street NW, Washington DC 20001 (www.national-academies.org/ilar).”

ORDER EULIPOTYPHLA

Sorex granarius (Lagranja Shrew/Long-Tailed Shrew)

2n=36

Perelman, Biltueva, and Graphodatsky (unpublished) Contributed by A. Graphodatsky

Sorex tundrensis (Tundra Shrew)

2n=30

Aniskin and Volobouev (1980)

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370

LAURASIATHERIA

Blarina carolinensis (Southern Short-Tailed Shrew)

2n=46

George et al. (1982) Contributed by R. J. Baker

Blarina carolinensis (Southern Short-Tailed Shrew)

2n=37

George et al. (1982) Contributed by R. J. Baker

ORDER EULIPOTYPHLA

Blarina carolinensis peninsulae (Southern Short-Tailed Shrew)

2n=50

George et al. (1982) Contributed by R. J. Baker

Blarina hylophaga (Elliot’s Short-Tailed Shrew)

2n=19

George et al. (1982) Contributed by R. J. Baker

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LAURASIATHERIA

Neomys fodiens (Old World Water Shrew)

2n=52

Perelman and Graphodatsky (unpublished) Contributed by A. Graphodatsky

ORDER EULIPOTYPHLA

Crocidura suaveolens (Lesser Shrew/White-Toothed Shrew)

2n=40

Perelman and Graphodatsky (unpublished) Contributed by A. Graphodatsky

Crocidura horsfieldii watasei (Horsfield’s Shrew/White-Toothed Shrew)

2n=26

Perelman and Graphodatsky (unpublished) Contributed by A. Graphodatsky

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374

LAURASIATHERIA

Crocidura pergrisea (Pale Gray Shrew/White-Toothed Shrew)

2n=22

A. Graphodatsky (unpublished)

Crocidura sibirica (Siberian Shrew/White-Toothed Shrew)

2n=40

A. Graphodatsky (unpublished)

ORDER EULIPOTYPHLA

Suncus murinus (Musk Shrew/Pygmy Shrew)

2n=40

Perelman, Biltueva, Graphodatsky, and Oda (unpublished) Contributed by A. Graphodatsky

2n=40

Perelman, Biltueva, Graphodatsky, and Oda (unpublished) Contributed by A. Graphodatsky

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376

LAURASIATHERIA

Crocidura dsinezumi (Dsinezumi Shrew/White-Toothed Shrew)

2n=40

Perelman, Biltueva, Graphodatsky, and Oda (unpublished) Contributed by A. Graphodatsky

Talpa europaea europaea (Old World Mole)

2n=34

Benirschke (1969)

ORDER EULIPOTYPHLA

Talpa altaica (Siberian Mole/Old World Mole)

2n=34

A. Graphodatsky (unpublished)

Parascalops breweri (Hairy-Tailed Mole)

2n=34

Benirschke (1969)

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LAURASIATHERIA

Order Chiroptera Approximately one-quarter (n = 977) of all mammal species are bats, so there are many species to be studied to document patterns and mechanisms associated with chromosomal change. Although bat karyotypic variation is generally described as conservative relative to other mammalian species, there are indeed many examples of extensive chromosomal rearrangement within bats. There is an unranked taxon, karyovarians (Baker et al., 2003), in the family Phyllostomidae (New World, leaf-nosed bats) that is named to recognize the extensive chromosomal evolution present in this clade, which includes 55 genera and 140 species. No member of this clade retains the proposed primitive karyotype for this complex of bats. In addition to the extreme morphological modifications required for flight (i.e., wings, 180° rotation of the hind limbs, etc.), bats possess a unique set of characteristics for a mammal of their body size. While there is considerable variation in the features listed, from an overview the following is true. Relative to other mammals of equal size and weight, bats have a longer life expectancy and generation time and a later gestation period. Several features of reproduction are also atypical. These include delayed implantation and embryonic diapause, delayed ovulation and fertilization, and sperm storage for several months (Neuweilier, 2000). Bats have a small litter size (one to four young per litter, but for most species one or two per pregnancy) and one or two litters per year. For many vespertilionids, a female will produce a single young per year. For microchiroptera there is a sophisticated development of echolocation and in megachiroptera the development of the visual system is advanced relative to other mammals. In both megachiroptera and microchiroptera, there is advanced encephalization of the brain (Stephan, 1977). Of course bats are more mobile than most other species of mammals, and some species migrate hundreds of miles annually. Greater mobility implies greater vagility; however, there is little scientific data available to understand how vagile bat species are. Further, the relative role of female versus male dispersal is poorly documented. Within bats there is a high level of diversity in social structuring. Sexual relationships characteristic of different species range from monogamy to polygamy and harems, in extreme to complete promiscuity, and in some cases what appears to be huge deme sizes (Neuweilier, 2000). Bats have a lower diploid number than is typical of other eutherian mammals. Diploid numbers range from 14 to 62 and the average has been reported to be 36.8 chromosomes (reviewed in Neuweilier, 2000). At the cellular level, the characteristics of the genome of bats are unique among mammals in that the total amount of DNA, the C-value, is less than is typical (50–87% of that of humans) (Burton et al., 1989). This reduction in genome size appears to be accomplished by a reduced amount of C-band positive material as well as a reduction in repetitive elements such as ribosomal genes and microsatellite clusters (Baker et al., 1979; Baker et al., 1992; Van Den Bussche et al., 1995). The karyotypic data from bats have been used in the literature for a number of studies with theoretical implications. An example is a study by Wilson et al. (1975), who used a review of bat karyotypic data, along with those from other mammals, to advance the idea that chromosomal evolution in mammals was driven by small deme sizes and inbreeding resulting from greater development of social structuring in mammals when compared to the social structure among amphibians, especially the order Anura. Wilson et al. (1975) extended their conclusions to include a cause-and-effect relationship between more chromosomal rearrangements and greater morphological evolution in mammals. Inbreeding as a consequence of social structuring has been questioned and rejected as a possible means of driving chromosomal change to fixation (Chesser and Baker, 1986). There are many examples where the karyotype has been radically reorganized without any substantial mor-

ORDER CHIROPTERA phological evolution, distinguishing the species with the radical reorganized karyotype from closely related species that have retained the primitive karyotype for the genus in question (Ellerman and Morrison-Scott, 1951; Fredga, 1977). Also, see the discussion below on karyotypic megaevolution. The canalization model of chromosomal evolution (Bickham and Baker, 1979) was described to explain the patterns of karyotypic change in bats and turtles. This model predicts that when an evolutionary lineage first enters a new grade of evolution (Simpson, 1948), the types of chromosomal rearrangements that will become established will be those that rearrange linkage groups. However, as morphology is canalized and becomes more adapted to this new grade, the types of chromosomal rearrangements that will become established will be those that are less disruptive to linkage groups (such as centric fusions and heterochromatic additions). The implications are that as the morphology becomes canalized, so will the karyotype, an example being when bats evolved from a terrestrial insectivore ancestor into a volant mammal. When bats first began to fly and exploit the volant grade, the karyotype was rearranged into linkage groups that more efficiently accommodated the developmental changes required to exploit this grade. Bats were used by Baker and Bickham (1980) to describe the phenomenon of karyotypic megaevolution. In karyotypic megaevolution, one species within a genus will have a radically reorganized karyotype such that it is difficult to relate the G-band patterns to those of congeneric species. Other species in the genus might exhibit little or no change from the primitive karyotypic condition associated with that genus. The first example of karyotypic megaevolution was described for the muntjacs (Ellerman and Morrison-Scott, 1951; Fredga, 1977). In this genus of barking deer, one species, Muntiacus reevesi (2n = 46), is thought to have an essentially unrearranged karyotype while M. muntjak greatly reorganized the karyotypes to produce 2n = 6 females and 2n = 7 males. The point here is that essentially all of the rearrangements that distinguish these two muntjak species have occurred in the 2n = 6, 7 species. Examples of this same pattern have been described for bats, including Lasionycteris noctivagans, Tonatia bidens, Tonatia schultzi, Micronycteris minuta, and Vampyress pusilla. Explaining how karyotypes in a one species can become so radically reorganized while closely related species experience little or no chromosomal evolution has been so difficult that these examples are a cytogenetic scandal. It has been argued that the number of chromosomal rearrangements could be used as a predictor of the number of population bottlenecks that a species had encountered during its evolution (Lande, 1979). However, there is no evidence that the species with radically reorganized karyotypes have social structuring or any other demographic factor that would disproportionately force them through numerous bottlenecks while the species with conserved karyotypes experienced none. These examples have been used to argue that the factors driving chromosomal evolution are not demographic but cellular, molecular, and meiotic (Baker et al., 1988). Karyotypes of the little yellow bats of the genus Rhogeessa (Genoways and Baker, 1996) were used to develop the model of speciation by monobrachial centric fusions (Baker and Bickham, 1986). The idea of differential fitness resulting from monobrachial homology involving centric fusions was introduced by Gropp et al. (1972) and further defined by Capanna (1982) for his emblematic model of chromosomal speciation. It has always been attractive to evolutionary biologists studying speciation to employ chromosomal rearrangements as a postmating isolating mechanism (King, 1993; White, 1975) and, L. H. Rieseberg (2001). This attraction primarily has been a product of chromosomal rearrangements reducing fertility (negative heterosis) in hybrids, which would help reduce gene flow between populations distinguished by chromosomal rearrangements (King, 1993). However, the paradox is: How can a chromosomal rearrangement become established in naturally

379

380

LAURASIATHERIA occurring populations if the rearrangement reduces heterozygosity sufficiently to be a postmating isolating mechanism? The idea that deme size and population bottlenecks were the solution to this dilemma was introduced by Wright (1941). Some cytogeneticists have gone to great lengths to solve this dilemma. For example, King (1993) suggested that meiotic drive could play a critical role in driving a new rearrangement to fixation in the face of negative heterosis in even a large population. Bickham (1995) pointed out that meiotic drive in King’s (1993) model must be transitory in nature and disappear after the fixation of the rearrangement in one population. Otherwise, the rearrangement will spread among all populations and not act as a reproductive isolating mechanism. Because this seems highly unlikely, the paradox of how to fix within a population a deleterious rearrangement that will subsequently perform well as a reproductive barrier has never been solved (Baker et al., 1988). The model of monobrachial speciation avoids this paradox because centric fusions are often population polymorphisms (Koop et al., 1983) and become fixed more easily in natural populations because new centric fusions do not result in unbalanced gamete formation to a point that they are highly selected against. In fact, it is not uncommon for many centric fusions to occur in a single species, as described by White (1975) as an example of karyotypic orthoselection. If different populations fix different centric fusions which have one arm that is homologous but the other arm attached to a different chromosome in one population and to a nonhomologous chromosome in another population, then a complex meiotic figure will result if hybridization occurs between these two populations (Baker and Bickham, 1986). The bottom line is that it is possible to establish chromosomal rearrangements independently in separate populations without significant selection against heterozygotes during the process of becoming fixed within that population. However, when these rearrangements undergo meiosis in a hybrid, the result is a much greater disadvantage (negative heterosis) to the hybrids than was encountered during the fixation process. Monobrachial speciation appears to be a very special model (does not occur in most species) that has been described for a variety of taxa, including the little yellow bat family Rhogeessa (Baker and Bickham, 1986; Dobigny et al., 2002; Kingswood et al., 2000; Qumsiyeh et al., 1999; Kumamoto et al., 1999; Gimenez et al., 1997; Searle, 1993; Piálek et al., 2001). In situ hybridization studies further document the potential for studies for Chiroptera karyotypes. Three papers (Volleth et al., 1999, 2001, 2002) clearly document the effectiveness of Zoo-FISH (fluorescence in situ hybridization) analysis in establishing syntenic groups within bats relative to humans. Further, these papers document the use of chromosomal evolution in establishing phylogenetic relationships. Parish et al. (2002), using in situ hybridization, document that these methodologies will be useful in understanding the relationship of genome organization, long interspersed nuclear elements (LINEs), and X chromosome inactivation. There is a tremendous amount of cytogenetic research remaining to be done within the order Chiroptera. Over half of the species of bats have yet to be described karyotypically even for diploid and fundamental numbers. In a field of science that is rapidly developing new methods of interrogating the karyotype, such as chromosomal paints, in situ hybridizations, genome sequences, and heterochromatin, few scientists are studying bat cytogenetics and genome organization. Less than 1% of the species of bats have been studied by modern techniques. Ah, so many bats, so many opportunities, and so little time. Note: Several contributors use Reference species in their numbering system. In the family Vespertilionidae the system established by J. B. Bickham uses Myotis nigricans. In the family Pyllostomidae the system established by Texas Tech uses Macrotys waterhousii. Robert J. Baker

ORDER CHIROPTERA

Rousettus aegyptiacus (Rousette Fruit Bat)

2n=36

Haiduk et al. (1981) Contributed by R. J. Baker

Rousettus leschenaulti (Rousette Fruit Bat)

2n=36

Sreepada and Gururaj (1995)

381

382

LAURASIATHERIA

Rousettus lanosus (Rousette Fruit Bat)

2n=36

Teeling and Stanyon (unpublished)

Rousettus (Lissonycteris) angolensis (Rousette Fruit Bat)

2n=36

Haiduk et al. (1981) Contributed by R. J. Baker

ORDER CHIROPTERA

Myonycteris torquata (Little Collared Fruit Bat)

2n=36

Haiduk et al. (1981) Contributed by R. J. Baker

Pteropus rodricensis (Rodriguez Flying Fox)

2n=38

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

383

384

LAURASIATHERIA

Pteropus giganteus (Indian Flying Fox)

2n=38

Choudhury and Jena (1995)

Pteropus lylei (Flying Fox)

2n=40

Hood et al. (1988) Contributed by R. J. Baker

ORDER CHIROPTERA

Hypsignathus monstrosus (Hammer-Headed Fruit Bat)

2n=36

Haiduk et al. (1981) Contributed by R. J. Baker

2n=36

Haiduk et al. (1981) Contributed by R. J. Baker

Epomops franqueti (Epauleted Bat)

385

386

LAURASIATHERIA

Micropteropus pusillus (Dwarf Epauleted Bat)

2n=35

Haiduk et al. (1981) Contributed by R. J. Baker

Scotonycteris ophiodon

2n=34

Haiduk et al. (1981) Contributed by R. J. Baker

ORDER CHIROPTERA

Otopteropus cartilagonodus (Long-Haired Tailless Flying Fox)

2n=48

Rickart et al. (1999)

Alionycteris paucidentata (Long-Haired Flying Fox)

2n=36

Rickart et al. (1999)

387

388

LAURASIATHERIA

Eonycteris robusta (Dawn Bat)

2n=36

Rickart et al. (1999)

Megaloglossus woermanni (African Long-Tongued Fruit Bat)

2n=36

Haiduk et al. (1981) Contributed by R. J. Baker

ORDER CHIROPTERA

Rhinopoma microphyllum (Mouse-Tailed Bat/Long-Tailed Bat)

2n=42

Qumsiyeh (1988)

Rhinopoma hardwickei (Mouse-Tailed Bat/Long-Tailed Bat)

2n=36

Qumsiyeh (1988)

389

390

LAURASIATHERIA

Taphozous longimanus (Tomb Bat)

2n=42

Sreepada et al. (1995)

Taphozous melanopogon (Tomb Bat)

2n=42

Sreepada et al. (1995)

Taphozous nudiventris (Tomb Bat)

2n=42

Sreepada et al. (1995)

ORDER CHIROPTERA

Taphozous saccolaimus (Tomb Bat)

2n=44

Sreepada et al. (1995)

Megaderma spasma (Asian False Vampire Bat)

2n=38

Morielle-Versute et al. (1992)

391

392

LAURASIATHERIA

Megaderma lyra (Asian False Vampire Bat)

2n=54

Choudhury and Mohanty (1993)

Rhinolophus rouxi (Horseshoe Bat)

2n=56

Zima et al. (1992) Contributed by M. Volleth

ORDER CHIROPTERA

Rhinolophus ferrumequinum (Horseshoe Bat)

2n=56

Teeling and Stanyon (unpublished)

393

394

LAURASIATHERIA

Hipposideros pomona (Old World Leaf-Nosed Bat)

2n=32

Sreepada et al. (1993)

Hipposideros ater (Old World Leaf-Nosed Bat)

2n=32

Sreepada et al. (1993)

Hipposideros fulvus (Old World Leaf-Nosed Bat)

2n=32

Sreepada et al. (1993)

ORDER CHIROPTERA

Hipposideros fulvus pallidus (Old World Leaf-Nosed Bat)

2n=32

Choudhury and Patro (1993)

Hipposideros cineraceus (Old World Leaf-Nosed Bat)

2n=32

Spreepada et al. (1993)

Hipposideros lankadiva (Old World Leaf-Nosed Bat)

2n=32

Spreepada et al. (1993)

395

396

LAURASIATHERIA

Hipposideros speoris (Old World Leaf-Nosed Bat)

2n=32

Spreepada et al. (1993)

ORDER CHIROPTERA

Ideogram: Pteronotus parnellii (Moustached Bat/Naked-Backed Bat/ Leaf-Lippped Bat)

2n=38

Sites et al. (1981) Contributed by J. W. Bickham

Pteronotus personatus (Moustached Bat/Naked-Backed Bat/ Leaf-Lippped Bat)

2n=38

Sites et al. (1981) Contributed by J. W. Bickham

397

398

LAURASIATHERIA

Pteronotus macleayii (Moustached Bat/Naked-Backed Bat/ Leaf-Lipped Bat)

2n=38

Sites et al. (1981) Contributed by J. W. Bickham

Pteronotus gymnonotus (Moustached Bat/Naked-Backed Bat/ Leaf-Lipped Bat)

2n=38

Sites et al. (1981) Contributed by J. W. Bickham

ORDER CHIROPTERA

Mormoops blainvillii (Leaf-Chinned Bat)

2n=38

Sites et al. (1981) Contributed by J. W. Bickham

Noctilio albiventris (Bulldog Bat/Fisherman’s Bat)

2n=34

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

399

400

LAURASIATHERIA

Desmodus rotundus (Common Vampire Bat)

2n=28

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

2n=28

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

ORDER CHIROPTERA

Diaemus youngi (White-Winged Vampire Bat)

2n=32

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

Diphylla ecaudata (Hairy-Legged Vampire Bat)

2n=32

Baker et al. (1979)

401

402

LAURASIATHERIA

Macrotus waterhousii (Big-Eared Bat)

2n=46

Baker et al. (1979)

Micronycteris megalotis (Little Big-Eared Bat)

2n=40

Baker et al. (1979)

ORDER CHIROPTERA

Micronycteris schmidtorum (Little Big-Eared Bat)

2n=38

Baker et al. (1979)

Micronycteris minuta (Little Big-Eared Bat)

2n=28

Baker et al. (1979)

403

404

LAURASIATHERIA

Micronycteris hirsuta (Little Big-Eared Bat)

2n=30

Baker et al. (1979)

Micronycteris brachyotis (Little Big-Eared Bat)

2n=32

Baker et al. (1979)

2n=28

Baker et al. (1979)

Micronycteris nicefori (Little Big-Eared Bat)

ORDER CHIROPTERA

Vampyrum spectrum (Linnaeus’s False Vampire Bat/Spectral Vampire Bat)

2n=30

Baker et al. (1979)

Trachops cirrhosus (Frog-Eating Bat)

2n=30

Baker et al. (1979)

405

406

LAURASIATHERIA

Chrotopterus auritus (Peter’s Woolly False Vampire Bat)

2n=28

Morielle-Versute et al. (1992)

Phylloderma stenops (Peter’s Spear-Nosed Bat)

2n=32

Baker et al. (1979)

ORDER CHIROPTERA

Phyllostomus discolor (Spear-Nosed Bat)

2n=32

Baker et al. (1979)

Phyllostomus hastatus (Spear-Nosed Bat)

2n=30

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

407

408

LAURASIATHERIA

Tonatia bidens (Round-Eared Bat)

2n=16

Baker et al. (1979)

Mimon crenulatum (Gray’s Spear-Nosed Bat)

2n=32

Baker et al. (1979)

ORDER CHIROPTERA

Glossophaga soricina (Long-Tongued Bat)

2n=32

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

2n=32

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

Monophyllus redmani (Leach’s Single-Leaf Bat)

2n=32

Baker et al. (1979)

409

410

LAURASIATHERIA

Lonchophylla robusta (Saussure’s Long-Nosed Bat)

2n=28

Baker et al. (1979)

Lonchophylla thomasi (Saussure’s Long-Nosed Bat)

2n=30

Baker et al. (1979)

2n=28

Baker et al. (1979)

Lionycteris spurrelli (Chestnut Long-Nosed Bat)

ORDER CHIROPTERA

Anoura caudifera (Geoffroy’s Long-Nosed Bat)

2n=30

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

2n=30

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

Anoura cultrata (Geoffroy’s Long-Nosed Bat)

2n=30

Baker et al. (1979)

411

412

LAURASIATHERIA

Lichonycteris obscura (Dark Long-Tongued Bat)

2n=24

Baker et al. (1979)

Hylonycteris underwoodi (Underwood’s Long-Tongued Bat)

2n=16

Baker et al. (1979)

Choeronycteris mexicana (Mexican Long-Nosed Bat/Hog-Nosed Bat)

2n=16

Baker et al. (1979)

ORDER CHIROPTERA

Choeroniscus intermedius (Godman’s Long-Nosed Bat)

2n=20

Stock (1975)

Erophylla sezekorni (Brown Flower Bat)

2n=32

Baker et al. (1979)

Choeroniscus gogmani (Godman’s Long-Nosed Bat)

2n=19

Hsu et al. (1968) Contributed by R. J. Baker

413

414

LAURASIATHERIA

Phyllonycteris poeyi (Cuban Flower Bat)

2n=32

Carollia castanea (Short-Tailed Leaf-Nosed Bat)

Baker et al. (1979)

2n=22

Stock (1975)

Carollia brevicauda (Short-Tailed Leaf-Nosed Bat)

2n=20F 2n=21M

Stock (1975)

ORDER CHIROPTERA

Carollia perspicillata (Short-Tailed Leaf-Nosed Bat)

2n=20(F) 21(M)

Teeling and Stanyon (unpublished)

Rhinophylla pumilio (Dwarf Little Fruit Bat)

2n=36

Baker et al. (1979)

415

416

LAURASIATHERIA

Rhinophylla fischerae (Fischer’s Little Fruit Bat)

2n=34

Sturnira lilium (Yellow Shoulder Bat/American Epauleted Bat)

Baker et al. (1979)

2n=30

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

2n=30

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

ORDER CHIROPTERA

Sturnira mordax (Yellow Shoulder Bat/American Epauleted Bat)

2n=30

Baker et al. (1979)

Sturnira erythromos (Yellow Shoulder Bat/American Epauleted Bat)

2n=30

Baker et al. (1979)

Uroderma magnirostrum (Tent-Building Bat)

2n=38

Baker et al. (1979)

417

418

LAURASIATHERIA

Platyrrhinus vittatus (White-Lined Bat)

2n=30

Baker et al. (1979)

Platyrrhinus vittatus lineatus (White-Lined Bat)

2n=30

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

2n=30

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

ORDER CHIROPTERA

Vampyressa pusilla (Yellow-Eared Bat)

2n=18

Baker et al. (1979)

Vampyressa nymphaea (Yellow-Eared Bat)

2n=26

Baker et al. (1979)

Vampyressa brocki (Yellow-Eared Bat)

2n=24

Baker et al. (1979)

419

420

LAURASIATHERIA

Chiroderma villosum (Big-Eyed Bat/White-Lined Bat)

2n=26

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

2n=26

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

ORDER CHIROPTERA

Chiroderma doriae (Big-Eyed Bat/White-Lined Bat)

2n=26

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

2n=26

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

421

422

LAURASIATHERIA

Mesophylla macconnelli (Little Yellow-Faced Bat)

2n=21

Baker et al. (1979)

Chiroderma improvisum (Big-Eyed Bat/White-Lined Bat)

2n=26

Baker et al. (1979)

Ectophylla alba (White Bat)

2n=30

Baker et al. (1979)

ORDER CHIROPTERA

Enchisthenes hartii (Hart’s Little Fruit Bat)

2n=31

Baker et al. (1979)

Artibeus jamaicensis (Neotropical Fruit Bat)

2n=30

Baker et al. (1979)

Ardops nichollsi (Tree Bat)

2n=31

Baker et al. (1979)

423

424

LAURASIATHERIA

Artibeus lituratus (Neotropical Fruit Bat)

2n=31

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

2n=31

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

ORDER CHIROPTERA

Phyllops haitiensis (Falcate-Winged Bat)

2n=31

Baker et al. (1979)

Ariteus flavescens (Jamaican Fig-Eating Bat)

2n=31

Baker et al. (1979)

2n=31

Baker et al. (1979)

Ametrida centurio (Little White-Shouldered Bat)

425

426

LAURASIATHERIA

Sphaeronycteris toxophyllum (Red Fruit Bat/Visored Bat)

2n=28

Baker et al. (1979)

Mystacina tuberculata (New Zealand Short-Tailed Bat)

2n=36

Bickham et al. (1980)

ORDER CHIROPTERA

Natalus major (Funnel-Eared Bat)

2n=36

Hoyt and Baker (1980) Contributed by R. J. Baker

Myotis keenii (Little Brown Bat)

2n=44

Bickham (1979)

Myotis thysanodes (Little Brown Bat)

2n=44

Bickham (1979)

427

428

LAURASIATHERIA

Myotis mystacina (Little Brown Bat)

2n=44

Volleth (1987)

Myotis sodalis (Little Brown Bat/Indiana Bat)

2n=44

Bickham (1979)

Myotis nigricans (Little Brown Bat)

2n=44

Bickham (1979)

ORDER CHIROPTERA

Ideogram: Myotis nigricans (Little Brown Bat)

2n=44

Bickham (1979)

Pipistrellus pipistrellus (Pipistrelle)

2n=44

Volleth (1987)

429

430

LAURASIATHERIA

Pipistrellus ceylonicus (Pipistrelle)

2n=36

Sreepada et al. (1996)

Pipistrellus javanicus (Pipistrelle)

2n=34

Volleth et al. (2001)

Pipistrellus coromandra (Pipistrelle)

2n=30

Sreepada et al. (1996)

ORDER CHIROPTERA

Pipistrellus (Hypsugo) crassulus (Pipistrelle)

2n=30

Volleth et al. (2001)

Pipistrellus kuhli (Pipistrelle)

2n=42

Volleth et al. (2001)

431

432

LAURASIATHERIA

Pipistrellus (Vespadelus) vulturnus (Pipistrelle)

2n=44

Volleth and Tidemann (1989)

Pipistrellus (Vespadelus) sagittula (Pipistrelle)

2n=44

Volleth and Tidemann (1989)

Pipistrellus stenopterus (Pipistrelle)

2n=32

Volleth et al. (2001)

ORDER CHIROPTERA

Pipistrellus (Falsistrellus) tasmaniensis (Pipistrelle)

2n=44

Volleth and Tidemann (1991)

2n=38

Volleth et al. (2001)

Pipistrellus mimus (Pipistrelle)

433

434

LAURASIATHERIA

Pipistrellus (Hypsugo) eisentrauti (Pipistrelle)

2n=42

Volleth et al. (2001)

Scotozous dormeri (Dormer’s Bat)

2n=30

Volleth et al. (2001)

ORDER CHIROPTERA

Hesperoptenus blanfordi (Blandford’s Bat)

2n=32

Chalinolobus morio (Chocolate Bat/Lobe-Lipped Bat/Groove-Lipped Bat/Wattled Bat)

Volleth et al. (2001)

2n=44

Volleth and Tidemann (1989)

Tylonycteris pachypus (Club-Footed Bat/Bamboo Bat)

2n=32

Volleth et al. (2001)

435

436

LAURASIATHERIA

Tylonycteris robustula (Club-Footed Bat/Bamboo Bat)

2n=46

Volleth et al. (2001)

Scotorepens balstoni (Lesser Broad-Nosed Bat)

2n=30

Volleth and Tidemann (1991)

ORDER CHIROPTERA

Eptesicus bottae (Big Brown Bat/House Bat/Serotine Bat)

2n=50

Volleth et al. (2001)

437

438

LAURASIATHERIA

Eptesicus fuscus (Big Brown Bat/House Bat/Serotine Bat)

2n=50

Teeling and Stanyon (unpublished)

Scotophilus kuhlii (House Bat/Yellow Bat)

2n=36

Stock (1975)

ORDER CHIROPTERA

Scotophilus heathi (House Bat/Yellow Bat)

2n=36

Stock (1975)

Corynorhinus townsendii (Lump-Nosed Bat/American Long-Eared Bat)

2n=32

Stock (1983)

439

440

LAURASIATHERIA

Lasiurus borealis (Hairy-Tailed Bat/Red Bat)

2n=28

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

2n=28

Varella-Garcia et al. (1989) Contributed by E. Morielle-Versute

Idionycteris phyllotis (Allen’s Big-Eared Bat/Lappet-Eared Bat)

2n=30

Stock (1983)

ORDER CHIROPTERA

Euderma maculatum (Spotted Bat/Pinto Bat)

2n=30

Stock (1983)

Murina cyclotis (Tube-Nosed Insectivora Bat)

2n=44

Rickart et al. (1999)

441

442

LAURASIATHERIA

Nyctophilus gouldi (Australian Big-Eared Bat)

2n=44

Volleth and Tidemann (1989)

Molossops temminckii (Broad-Faced Bat)

2n=48

Morielle-Versute et al. (1996)

ORDER CHIROPTERA

Molossops abrasus (Broad-Faced Bat)

2n=34

Morielle-Versute et al. (1996)

Nyctinomops laticaudatus (New World Free-Tailed Bat)

2n=48

Morielle-Versute et al. (1996)

Eumops perotis (Mastiff Bat/Bonneted Bat)

2n=48

Morielle-Versute et al. (1996)

443

444

LAURASIATHERIA

Eumops glaucinus (Mastiff Bat/Bonneted Bat)

2n=40

Morielle-Versute et al. (1996)

Molossus molossus (Velvety Free-Tailed Bat)

2n=48

Morielle-Versute et al. (1996)

Molossus rufus (ater) (Velvety Free-Tailed Bat)

2n=48

Morielle-Versute et al. (1996)

ORDER CARNIVORA

Order Carnivora Synthesis of Carnivore Chromosome Evolution Most authorities recognize two major phylogenetic clades within Carnivora, caniformia and feliformia. Although the systematics of carnivores is currently in a state of revision, we are with three exceptions, following the format of Ronald Nowack (1999). (1) We include Pinnipedia in the caniformia branch of carnivores and (2) we use the phylogenetic classification of Johnson et al. (1997) for Felidae (3) recognize the family Nandiniidae as being basal to all other feliformia (McKenna and Bell, 1997; Flynn et al., 2005). Caniformia consists of seven families: Canidae, Ursidae, Procyonidae, Mustelidae, Otariidae, Odobenidae, and Phocidae. Feliformia consists of five families: Nandiniidae, Viverridae, Herpestidae, Hyaenidae, and Felidae. This order of dogs, bears, raccoons, weasels, seals, civets, mongooses, hyenas, and cats consists of 11 families, 115 genera, and 280 species with a worldwide distribution. A large body of molecular data, including Zoo-FISH, which demonstrates chromosome homology between species by DNA hybridization, has shown that the euchromatic portions of the genomes of all carnivores studied thus far are equivalent. This information in combination with high-quality Gbanding has demonstrated that to a very large extent euchromatic G-bands have been neither lost nor gained during carnivore chromosome evolution. The evidence for this statement is the following. When rearrangements that differentiate one species from another as defined by Zoo-FISH and G-banding are reversed, what results are two karyotypes that look the same. The heterochromatic portions of the genomes, on the other hand, have evolved quite independently in different carnivore species. In the early 1970s, Doris Wurster-Hill began defining ancestral carnivore chromosomes on the basis of G-banding alone. Chromosomes with the same G-banding patterns that were found widespread in families on both branches of the carnivore lineage, which split early in the radiation, were deemed ancestral. The description of ancestral chromosomes in the last 30 years has been refined and confirmed by a number of independent investigators, particularly with extensive Zoo-FISH analysis. The ancestral carnivore karyotype (ACK) likely consists of 2n = 38 chromosomes, most of which are conserved as full chromosomes or chromosome arms in the majority of carnivore species in all families except Ursidae and Canidae (Nash et al., 1998, 2001; Nie et al., 2002; Perelman et al., 2005; Nash et al., 2006). The pattern of chromosome evolution that shaped the karyotypes of all modern carnivores proceeded along one of two distinctive pathways and correlated perfectly with family lineage. Only Ursidae and Canidae experienced a dramatic increase in chromosome number early in their family lineages: from 2n = 38 to 2n = 72 in Ursidae and 2n = 38 to 2n = ~78 in Canidae. As a result modern ursid and canid karyotypes are distinct from other carnivores. Modern species of all other carnivore families are derived from the original low-number ACK, 2n = 38. A majority of the arm associations found in the 15 biarmed chromosomes of the ACK are preserved in all species, and other than a few inversions, all the G-banding patterns of the euchromatic arms are preserved as well. As a result their karyotypes are all similar. With regard to change in the chromosome number, carnivore karyotypes evolved in a Robertsonian fashion. Centric fissions increased chromosome number while centric fusions reduced chromosome number. In karyotypes with very low chromosome numbers, such as the pinnipeds, tandem fusions are increasingly common. In karyotypes with very high chromosome numbers, chromosome arm fragmentation occured and did so in a two-step fashion. The first step involves one of the most common rearrangements in

445

446

LAURASIATHERIA mammalian chromosome evolution, an inversion which changes an acrocentric to a biarmed chromosome. A centric fission of this derived chromosome then resulted in two chromosomes derived from one ancestral acrocentric chromosome. Centric fissions and chromosome arm fragmentations were responsible for the evolution of the high-number ancestral ursid karyotype (AUK) from the ACK. We can state this with confidence because the chromosomes resulting from these centric fissions and fragmentations are found largely unchanged in the chromosomes of modern bear karyotypes. Evolution of the ancestral karyotype of extant canids (AKEC) from the ACK was similar but more complicated than in Ursidae and probably involved an initial expansion, a reduction, and a second expansion of chromosome number. As a result many dog chromosomes are mosaics of two to four ancestral arm fragments (Nash et al., 2001) Although the karyotypes of ursids and canids have diverged significantly from the ACK, the chromosomes of species within each family are highly conserved, as are other carnivore families. When related carnivore species do differ significantly in the amount of chromosomal material present in their karyotypes, it is due to the addition of species-specific constitutive heterochromatin. This heterochromatin may result in de novo short or long arms, blocks of pericentric or terminal material, or occasionally interstitial insertions. When interstitial insertions occur, they may change the G-banded gestalt of the chromosome.

Family Canidae The family Canidae includes 36 living species that are divided into 16 genera. The G-banded karyotypes of 14 species from 12 genera are present in the atlas. Zoo-FISH and G-banding analysis demonstrate that most of the chromosome arms of the low-number canid species show G-banding homology to acrocentric chromosomes of species with high-number karyotypes. When the biarmed chromosomes of low-number species are compared to each other, the chromosome arms are joined in different combinations in different species (Wayne et al., 1987a,b; Nash et al., 2001). These observations suggest that the low-number karyotype canids evolved from a common high-number ancestral karyotype mainly through a series of independent centric fusions. A second independent observation strongly supports this hypothesis. Most of the ACK chromosome arm breakpoints resulting from fragmentation are the same in both high- and low-chromosome-number canids karyotypes. Since chromosome arm fragmentation in carnivores only occurs in karyotypes experiencing a concerted increase in chromosome number, the high-number karyotypes of canids must be ancestral to the low-number karyotypes. The evolution of the AKEC (2n = ~78) from the ACK (2n = 38) cannot be explained as a single-step process as demonstrated in Ursidae. The 10 centric fissions and 30 chromosome arm fragmentations that separate the two karyotypes would result in an AKEC with 2n = 122 chromosomes. Probably two separate expansions of chromosome number occurred between the ACK and AKEC. During the first expansion, approximately half of the 30 ACK arm breaks found in the AKEC may have occurred, resulting in a few simple mosaic chromosomes (consisting of only two ancestral arm fragments) and would appear as the AUK. If the first expansion was followed by a concerted decrease to a low-number karyotype, as found in the red fox Vulpes vulpes (VVU, 2n = 32), for example, then tandem fusions would have been favored resulting in more mosaic chromosomes. A second expansion of the chromosome number would produce the additional ACK chromosome arm breakpoints and the chromosome number found in the AKEC. Pericentric and

ORDER CARNIVORA paracentric inversions during this period of chromosome evolution would further increase the number and complexity (up to four ACK arm fragments per chromosome) of the mosaic chromosomes, characteristic of the AKEC and all modern dog karyotypes.

Family Ursidae The family Ursidae consists of three genera and eight species. G-banded karyotypes of all eight species are presented (O’Brien et al., 1985; Nash and O’Brien, 1987; Nash et al., 2001). The three bear genera each have a different karyotype. Six ursine bear species share a nearly identical G-banded karyotype consisting of mostly acrocentric chromosomes (2n = 74). The spectacled bear Tremarctos ornatus (TOR), which diverged from the ursine bears some 6 mya, has 2n = 52 largely biarmed chromosomes. G-banding and Zoo-FISH analysis reveals that the reduction of chromosome number in the spectacled bear resulted from 11 centric fusions of acrocentric ursine bear chromosomes. The G-banding patterns of all the homologous chromosome arms of the spectacled bear and ursine bears are the same. The giant panda Ailuropoda melanoleuca (AME), which diverged from the ancestral ursid lineage some 12 mya, consists of 2n = 42 chromosomes, of which all but three are biarmed. Chromosome number in the giant panda has been reduced as the result of 13 centric fusions and 3 tandem fusions. The giant panda shares eight of nine ursid specific ACK chromosome arm breakpoints confirming its unambiguous membership in the bear family. The evolution of the AUK from the ACK can be viewed as a single continuous process. It involved eight centric fissions, eight chromosome arm fragmentations, and two subsequent tandem fusions of chromosome arm fragments of the ACK. These three kinds of chromosomal rearrangements alone gave rise to the 2n = 72 AUK. Thus, new chromosome arm associations and cytogenetically observable inversions of ACK chromosomes played a minor role in shaping the AUK chromosomes. This simple scenario is confirmed by the observation that the ACK chromosome G-banding patterns are conserved in all modern ursid chromosomes.

Family Procyonidae The family Procyonidae has 19 species divided into 2 subfamilies and 7 genera. The G-banded karyotypes of 5 species are presented and represent species from both subfamilies. The 2n chromosome number is 38 except for the lesser panda Ailurus fulgens (AFU), which has 2n = 36. Zoo-FISH using domestic cat painting probes have been performed on the lesser panda, the raccoon Procyon lotor (PLO), and the ringtail Bassariscus astutus (BAS). The hybridization patterns of the raccoon and ringtail are identical and their G-banding patterns are the same as well. By G-banding alone the kinkajou Potos flavus (PFL) chromosomes are also the same as those of PLO and BAS. The ringtail karyotype has the same chromosome number as the ACK 2n = 38. G-banding reveals a single pericentric inversion that distinguishes the ringtail, raccoon, and kinkajou from the ACK. The lesser panda has a tandem fusion of an ACK biarmed and acrocentric chromosome (2n = 36). Five chromosomes of the lesser panda differ from their ACK homologues as a result of pericentric inversions.

Family Mustelidae The family Mustelidae consists of 67 species divided into 5 subfamilies and 25 genera. The G-banded karyotypes of 26 species from 11 genera are presented and represent species from all but one subfamily. The 2n chromosome

447

448

LAURASIATHERIA number ranges from 30 to 44, although 20 of these species have 2n = 38. ZooFISH using domestic cat painting probes has been performed on 9 species representing 3 subfamilies and 5 genera. Some species have been painted with human, domestic dog, raccoon dog, and giant panda probes as well (Graphodatsky et al., 2000a,b; Nie et al., 2002). Zoo-FISH and G-banding analysis reveal that mustelid karyotypes are highly conserved relative to the ACK. As in most other carnivore families, Robertsonian chromosome evolution is the major force shaping mustelid karyotypes. The ACK chromosome arms are not broken but are simply reshuffled by centric fissions and fusions. In carnivores, the incidence of tandem fusion increases in low-number karyotypes because the ACK has only six acrocentric chromosomes that can reduce the chromosome number through centric fusions. The American mink Mustela visons (MVI) has 2n = 30 chromosomes. This reduction in chromosome number from the ACK is the result of two centric fissions, one centric and five tandem fusions. Four of these tandem fusions are between a biarmed and an acrocentric chromosome, and one involves two biarmed chromosomes. As a result six ancestral centromeres are inactivated. Zoo-FISH and Gbanding analysis reveal that two pericentric and five paracentric inversions separate the ACK and MVI karyotypes as well.

Families Otariidae, Odobenidae, and Phocidae The karyotypes of species in these three families are highly conserved and will be described together. The family Otariidae consists of 14 species and 7 genera. G-banded karyotypes of four species from four genera are presented. The 2n chromosome number is 36 for all four species. The family Odobenidae is monotypic, with the walrus Odobenis rosmarus (ORO) being its only living representative. Its G-banded karyotype is presented (2n = 32). The family Phocidae consists of 19 species and 10 genera. G-banded karyotypes of 10 species from 7 genera are presented. The 2n chromosome number is either 32 or 34 depending on the presence or absence of a centric fusion. Zoo-FISH using human painting probes has been performed on one species, the harbor seal Phoca vitulina (PVI), family Phocidae (Frönicke et al., 1997). Because the chromosome segment homologues between human and cat are well known, one can easily convert from human to domestic cat to ACK chromosomes for comparison to the harbor seal. The harbor seal karyotype differs from the ACK by three tandem fusions, and one pericentric inversion, 2n = 34. The G-banded karyotypes of all phocids appear to be the same with the exception that 2n = 34 species have the homologues to the domestic cat chromosomes E1 and A1q separate, while 2n = 32 species have these chromosomes fused. Based on G-banding analysis the karyotypes of otariid species differ from the 2n = 34 phocids by the absence of one tandem fusion. The homologues of domestic cat chromosomes F1 and D2 are fused in phocids and separate in otariids. Also, the homologue of domestic cat chromosome E3, which is tandemly fused to the homologue of domestic cat chromosome B4 in phocids, is instead tandemly fused to the homologue of domestic cat chromosome F3 in otariids. These latter two chromosome arrangements are also found in the walrus karyotype. While most of the walrus chromosomes are the same as other pinnipeds, a few differences will require Zoo-FISH analysis to resolve.

Family Nandiniidae The family Nandiniidae consists of one living representative, Nandinia binotata (NBI), which is basal to all other feliformia (McKenna and Bell, 1997; Flynn et al., 2005). Its karyotype 2n = 38, is very similar to the ringtail (BAS).

ORDER CARNIVORA Fifteen of 18 autosomes in the two species appear identical morphologically and by their longitudinal banding patterns. It is the only feliform species to have the three signature caniform chromosomes, as represented by ringtail (BAS) chromosomes 1, 3, and 4, in its karyotype.

Family Viverridae The family Viverridae consists of 34 species divided into four subfamilies and 18 genera. The G-banded karyotypes of nine species from eight genera are presented and represent at least one species from each subfamily. The 2n chromosome number ranges from 34 to 54, although the most common chromosome number is 2n = 42. Zoo-FISH using domestic cat painting probes has been performed on two viverrid species the masked palm civet Paguma larvata (PLA) 2n = 44, and the Malagasy civet Fossa fossana (FFO) 2n = 44. The masked palm civet karyotype differs from the ACK by four centric fissions and one tandem fusion (Perelman et al., 2005). Based on G-banding analysis alone, the ACK chromosomes or chromosome arms in the better banded viverrid karyotypes are easily recognizable. In general, viverrid karyotypes differ from one another and the ACK by a few centric fissions and fusions resulting in some new arm associations. Paracentric and pericentric inversions also occur, but tandem fusions are rare. The ACK chromosome arms remain intact with the possible exception of two species of genets, the small spotted genet Genetta genetta (GGE) and Genetta tigrina (GTI), both of which have fairly high chromosome number karyotypes 2n = 54. While most ACK chromosomes or chromosome arms are intact, a few small chromosomes appear to be fragments of ACK chromosome arms. This observation reinforces the idea that breakage of ACK chromosome arms in carnivore species only occurs when a karyotype experiences a concerted increase in chromosome number. In this example ACK chromosome arm fragmentation occurred late in the radiation of modern viverrids. In Ursidae and Canidae ACK chromosome arm fragmentation occurred before the radiation of modern bears and dogs.

Family Herpestidae The family Herpestidae has 19 genera and 39 species grouped into three subfamilies. The G-banded karyotypes of six species from six genera are presented and represent at least one species from each subfamily. The 2n chromosome number ranges from 32 to 44. Zoo-FISH using domestic cat painting probes has been performed on one herpestid species, the dwarf mongoose Helgale pavula (HPA). The dwarf mongoose karyotype 2n = 36 differs from the ACK karyotype by four centric fissions, three centric fusions, two tandem fusions, and one pericentric inversion. Several of the centric fission chromosome arms were subsequently involved in centric fusions giving rise to new chromosome arm associations. The two tandem fusions each involved a biarmed and acrocentric chromosome. The centromeres of the biarmed chromosomes were inactivated. The pericentric inversion changed an acrocentric to a biarmed chromosome. No ACK chromosome arms have been fragmented in the dwarf mongoose karyotype, and this appears to be the case in the other herpestids based on their G-banded chromosomes.

Family Hyaenidae The family Hyaenidae has four genera and four species. G-banded karyotypes of the brown hyena Parahyaena brunnea (PHR) and the spotted hyena Crocuta

449

450

LAURASIATHERIA crocuta (CCR) are presented. The 2n chromosome number for both species is 40. Zoo-FISH using human, American mink (MVI), and domestic dog (Canis familiaris, CFA) painting probes has been performed on the spotted hyena (Perelman et al., 2005). The spotted hyena karyotype differs from the ACK by six centric fissions, three centric fusions, one combination of a centric fusion plus tandem fusion, and six pericentric inversions. The 23 autosomal chromosome arms of the hyena not involved in inversions have the same G-banding pattern as their ACK chromosome arm homologues. All ACK chromosome arms remain intact (i.e., not fragmented) in the spotted hyena. Even though a different numbering system is used in the brown hyena karyotype, its chromosomes appear to have a one-for-one G-banding homology to the spotted hyena chromosomes.

Family Felidae The family Felidae consists of 17 genera and 38 species. G-banded karyotypes exist for every species except Oreailarus jacobita (OJA). The karyotypes of 18 species of felids are presented in this atlas. The chromosome number of Felidae species is 2n = 38, with the exception of the South American felids (2n = 36). Chromosomes of all Felidae species are so similar that a single karyotype format is used for all felids (Wurster-Hill and Gray, 1975; Wurster-Hill and Centerwall, 1982). The only obvious chromosomal difference between Felidae species involves two small acrocentric chromosomes F2 and F3; F3 is the ancestral version of domestic cat (Felis Catus, FCA) chromosome F1 and is found in most felid species. An example of each F2 and F3 chromosome variant is described below. Chromosomes F2 and F3 are fused at their centromeres to form the metacentric chromosome C3 in the South American cats. In several other felids, chromosome F3 is missing and is replaced by the small metacentric chromosome E4. Chromosome E4 is the result of a pericentric inversion in F3. In Herpallarua yagouaroudi (HYA) both F2 and F3 are similarly inverted to form chromosomes E4 and E5. The other chromosome changes consist of a handful of subtle inversions and small additions of C-banded material (heterochromatin) in a few chromosomes of some species. Zoo-FISH between the domestic cat and species of other carnivore families indicates that most felid chromosomes have a one-for-one homology with ACK chromosomes. The domestic cat karyotype differs from the ACK by three centric fissions, three centric fusions, one large pericentric inversion, and the formation of a neocentromere in chromosome C2. Reciprocal chromosome painting between the domestic cat and species from other mammalian orders (human, pig) demonstrates that large linkage groups are preserved between orders as well. William G. Nash

ORDER CARNIVORA

Vulpes vulpes (Red Fox)

2n=34+B’s

Graphodatsky and Beklemisheva (unpublished) Contributed by A. Graphodatsky

451

452

LAURASIATHERIA

Ideogram: Vulpes vulpes (Red Fox)

2n=34+B’s

Rubtsova (1998)* * “Reprinted with permission from ILAR Journal, 39: 182–188 (1998), Institute for Laboratory Animal Research, The Keck Center of the National Academies, 500 Fifth Street NW, Washington DC 20001 (www.national-academies.org/ilar).”

ORDER CARNIVORA

Vulpes corsac (Corsac Fox)

2n=36

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Vulpes macrotis (Kit Fox)

2n=50

Wayne et al. (1987a)

453

454

LAURASIATHERIA

Fennecus zerda (Fennec Fox)

2n=64

Graphodatsky (unpublished)

ORDER CARNIVORA

Alopex lagopus (Arctic Fox)

2n=50

Nash et al. (2001)

455

456

LAURASIATHERIA

Ideogram: Alopex lagopus (Arctic Fox)

2n=50

Nash et al. (2001)

ORDER CARNIVORA

Urocyon cinereoargenteus (Grey Fox)

2n=66

Wayne et al. (1987a)

457

458

LAURASIATHERIA

Cerdocyon thous (Crab-Eating Fox)

2n=74

Nash et al. (2001)

ORDER CARNIVORA

Ideogram: Cerdocyon thous (Crab-Eating Fox)

2n=74

Nash et al. (2001)

459

460

LAURASIATHERIA

Nyctereutes procyonoides viverrinus (Japanese Raccoon Dog)

2n=38 + B’s

Nash et al. (2001)

ORDER CARNIVORA

Ideogram: Nyctereutes procyonoides viverrinus (Japanese Raccoon Dog)

2n=38 + B’s

Nash et al. (2001)

461

462

LAURASIATHERIA

Nyctereutes procyonoides procyonoides (Chinese Raccoon Dog)

2n=54+B’s

Graphodatsky, Beklemisheva and Yang (unpublished) Contributed by A. Graphodatsky

ORDER CARNIVORA

Atelocynus microtis (Small-Eared Dog/Round-Eared Dog)

2n=74

Wurster-Hill and Centerwall (1982)

463

464

LAURASIATHERIA

Speothos venaticus (Bush Dog)

2n=74

Wayne et al. (1987a)

ORDER CARNIVORA

Canis latrans (Coyote)

2n=78

Wurster-Hill and Centerwall (1982)

465

466

LAURASIATHERIA

Canis lupus (Grey Wolf)

2n=78

Wurster-Hill and Centerwall (1982)

ORDER CARNIVORA

Canis familiaris (Domestic Dog)

2n=78

Graphodatsky et al. (2000a)

467

468

LAURASIATHERIA

Ideogram: Canis familiaris (Domestic Dog)

2n=78

Graphodatsky et al. (2000b)

ORDER CARNIVORA

Chrysocyon brachyurus (Maned Wolf)

2n=76

Wayne et al. (1987a)

469

470

LAURASIATHERIA

Otocyon megalotis (Bat-Eared Fox)

2n=72

Wayne et al. (1987a)

ORDER CARNIVORA

Tremarctos ornatus (Spectacled Bear)

2n=52

Nash (unpublished)

471

472

LAURASIATHERIA

Ideogram: Tremarctos ornatus (Spectacled Bear)

2n=52

Nash (unpublished)

ORDER CARNIVORA

Ursus thibetanus (Asiatic Black Bear)

2n=74

Nash and O’Brien (1987)

473

474

LAURASIATHERIA

Ursus americanus (American Black Bear)

2n=74

Nash and O’Brien (1987)

ORDER CARNIVORA

Ursus arctos (Brown Bear)

2n=74

Nash and O’Brien (1987)

475

476

LAURASIATHERIA

Ursus maritimus (Polar Bear)

2n=74

Nash and O’Brien (1987)

ORDER CARNIVORA

Ursus malayanus (Sun Bear)

2n=74

Nash and O’Brien (1987)

477

478

LAURASIATHERIA

Ursus ursinus (Sloth Bear)

2n=74

Nash and O’Brien (1987)

ORDER CARNIVORA

Ailuropoda melanoleuca (Giant Panda)

2n=42

Nash (unpublished)

479

480

LAURASIATHERIA

Ideogram: Ailuropoda melanoleuca (Giant Panda)

2n=42

Nash (unpublished)

ORDER CARNIVORA

Ailurus fulgens (Lesser Panda/Red Panda)

2n=36

Nash (unpublished)

2n=38

Nash (unpublished)

Bassariscus astutus (Ringtail/Cacomistle)

481

482

LAURASIATHERIA

Potos flavus (Kinkajou)

2n=38

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Procyon lotor (Raccoon)

2n=38

Stanyon (unpublished)

ORDER CARNIVORA

Ideogram: Procyon lotor (Raccoon)

2n=38

Stanyon et al. (1993)

483

484

LAURASIATHERIA

Bassaricyon sp. (Olingo)

2n=38

Wurster-Hill and Gray (1975)

Mustela erminea (Ermine/Stoat)

2n=44

Graphodatsky (unpublished)

ORDER CARNIVORA

Mustela nivalis (Least Weasel)

2n=42

2n=42

Perelman and Graphodatsky (unpublished)

Perelman and Graphodatsky (unpublished)

Contributed by A. Graphodatsky

Contributed by A. Graphodatsky

485

486

LAURASIATHERIA

Mustela altaica (Mountain Weasel)

2n=44

2n=44

Perelman and Graphodatsky (unpublished)

Perelman and Graphodatsky (unpublished)

Contributed by A. Graphodatsky

Contributed by A. Graphodatsky

ORDER CARNIVORA

Mustela lutreola (European Mink)

2n=38

Graphodatsky (unpublished)

2n=38

Graphodatsky (unpublished)

Mustela sibirica (Siberian Weasel)

487

488

LAURASIATHERIA

Mustela sibirica itatsi (Japanese Weasel)

2n=38

Graphodatsky (unpublished)

2n=30

Nash (unpublished)

Mustela vison (American Mink)

ORDER CARNIVORA

489

Ideogram: Mustela vison (American mink)

2n=30

Serov (1998)*

Mustela putorius putorius (Wild Polecat)

2n=40

2n=40

Graphodatsky (unpublished)

Graphodatsky (unpublished)

* “Reprinted with permission from ILAR Journal, 39: 189–194 (1998), Institute for Laboratory Animal Research, The Keck Center of the National Academies, 500 Fifth Street NW, Washington DC 20001 (www.national-academies.org/ilar).”

490

LAURASIATHERIA

Mustela putorius furo (Domesticated Ferret)

2n=40

2n=40

Graphodatsky (unpublished)

Graphodatsky (unpublished)

ORDER CARNIVORA

Mustela eversmanni (Steppe Polecat)

2n=38

Graphodatsky (unpublished)

Martes foina (Beach Marten)

2n=38

Nie et al. (2002) Contributed by F. Yang

491

492

LAURASIATHERIA

Vormela peregusna (Marbled Polecat)

2n=38

2n=38

Graphodatsky (unpublished)

Graphodatsky (unpublished)

ORDER CARNIVORA

Martes martes (Pine Marten)

2n=38

Graphodatsky (unpublished)

2n=38

Graphodatsky (unpublished)

Martes zibellina (Sable)

493

494

LAURASIATHERIA

Martes melampus (Japanese Marten)

2n=38

Perelman, Graphodatsky, and Robinson (unpublished) Contributed by A. Graphodatsky

Martes flavigula (Yellow-Throated Marten)

2n=40

Nie et al. (2002) Contributed by F. Yang

ORDER CARNIVORA

Eira barbara (Tayra)

2n=38

Wurster-Hill and Centerwall (1982)

Galictis vittata (Allamand’s Grisón/Greater Grisón)

2n=38

Wurster-Hill and Centerwall (1982)

495

496

LAURASIATHERIA

Ictonyx striatus (Striped Polecat/Zorilla)

2n=38

2n=38

Perelman, Graphodatsky, and Robinson (unpublished)

Perelman, Graphodatsky, and Robinson (unpublished)

Contributed by A. Graphodatsky

Contributed by A. Graphodatsky

ORDER CARNIVORA

Gulo gulo (Wolverine)

2n=42

Graphodatsky (unpublished)

Mellivora capensis (Ratel/Honey Badger)

2n=40

Wurster-Hill and Centerwall (1982)

497

498

LAURASIATHERIA

Meles meles (Old World Badger)

2n=44

Nie et al. (2002) Contributed by F. Yang

Melogale sp. (Ferret Badger)

2n=38

Wurster-Hill and Centerwall (1982)

ORDER CARNIVORA

Spilogale gracilis latifrons (Spotted Skunk)

2n=60

Benirschke (1969)

Spilogale putorius interrupta (Spotted Skunk)

2n=64

Benirschke (1969)

499

500

LAURASIATHERIA

Lutra lutra (European River Otter)

2n=36

Graphodatsky (unpublished)

Nandinia binotata (African Palm Civet)

2n=38

Wurster-Hill and Gray (1975)

ORDER CARNIVORA

Genetta genetta (Small Spotted Genet)

2n=54

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Genetta tigrina (Large Spotted Genet)

2n=54

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

501

502

LAURASIATHERIA

Prionodon linsang (Banded Linsang/Oriental Linsang)

2n=34

Wurster-Hill and Gray (1975)

Viverricula indica (Lesser Oriental Civet/Rasse)

2n=38

Wurster-Hill and Gray (1975)

ORDER CARNIVORA

Paradoxurus hermaphroditus (Palm Civet)

2n=42

Wurster-Hill and Gray (1975)

Arctictis binturong (Binturong)

2n=42

Wurster-Hill and Gray (1975)

503

504

LAURASIATHERIA

Paguma larvata (Masked Palm Civet)

2n=44

Graphodatsky and Beklemisheva (unpublished) Contributed by A. Graphodatsky

ORDER CARNIVORA

Hemigalus derbyanus (Banded Palm Civet)

2n=42

Wurster-Hill and Gray (1975)

Fossa fossana (Malagasy Civet)

2n=42

Nash (unpublished)

505

506

LAURASIATHERIA

Galidia elegans (Malagasy Ring-Tailed Mongoose)

2n=44

Wurster-Hill and Gray (1975)

Herpestes javanicus (Java Mongoose)

2n=36

Yang and Nie (unpublished) Contributed by N. W. Nie

ORDER CARNIVORA

Helogale parvula (Dwarf Mongoose)

2n=36

Nash (unpublished)

Bdeogale sp. (Black-Legged Mongoose)

2n=36

Wurster-Hill and Gray (1975)

507

508

LAURASIATHERIA

Atilax paludinosus (African Marsh Mongoose/Water Mongoose)

2n=35

Wurster-Hill and Gray (1975)

Cryptoprocta ferox (Fossa)

2n=42

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CARNIVORA

Parahyaena brunnea (Brown Hyena)

2n=40

Perelman and Graphodatsky (unpublished) Contributed by A. Graphodatsky

509

510

LAURASIATHERIA

Crocuta crocuta (Spotted Hyena)

2n=40

Nash (unpublished)

ORDER CARNIVORA

Leopardus pardalis (Ocelot)

2n=36

Wurster-Hill and Gray (1975)

Leopardus tigrinus (Tiger Cat/Little Spotted Cat)

2n=38

Wurster-Hill and Centerwall (1982)

511

512

LAURASIATHERIA

Lynchailurus colocolo (Pampas Cat)

2n=38

Wurster-Hill and Gray (1973)

ORDER CARNIVORA

Oncifelis geoffroyi (Geoffroy’s Cat)

2n=36

Nash (unpublished)

513

514

LAURASIATHERIA

Felis catus (Domestic Cat)

2n=38

Nash (unpublished)

ORDER CARNIVORA

Ideogram: Felis catus (Domestic Cat)

2n=38 Cho et al. (1997)

Contributed by H. Satoh

515

516

LAURASIATHERIA

Felis chaus (Jungle Cat)

2n=38

Wurster-Hill and Gray (1973)

Felis margarita (Sand Cat)

2n=38

Wurster-Hill and Gray (1975)

ORDER CARNIVORA

Felis nigripes (Black-Footed Cat)

2n=38

Nash (unpublished)

517

518

LAURASIATHERIA

Panthera leo (Lion)

2n=38

Nash (unpublished)

ORDER CARNIVORA

Panthera tigris altaica (Siberian Tiger)

2n=38

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

519

520

LAURASIATHERIA

Neofelis nebulosa diardi (Bornean Clouded Leopard)

2n=38

Nash (unpublished)

ORDER CARNIVORA

Neofelis nebulosa (Mainland Clouded Leopard)

2n=38

Nash (unpublished)

521

522

LAURASIATHERIA

Puma concolor (Puma/Cougar/Mountain Lion)

2n=38

Wurster-Hill and Gray (1973)

ORDER CARNIVORA

Acinonyx jubatus (Cheetah)

2n=38

Nash (unpublished)

523

524

LAURASIATHERIA

Herpailurus yagouaroundi (Jaguarundi)

2n=38

Wurster-Hill and Gray (1973)

2n=38

Yang and Li (unpublished) Contributed by T. Li

Lynx lynx (Eurasian Lynx)

ORDER CARNIVORA

Prionailurus bengalensis (Asian Leopard Cat)

2n=38

Wurster-Hill and Gray (1973)

525

526

LAURASIATHERIA

Prionailurus viverrinus (Fishing Cat)

2n=38

Wurster-Hill and Gray (1973)

ORDER CARNIVORA

Caracal caracal (Caracal)

2n=38

Wurster-Hill and Gray (1973)

2n=38

Wurster-Hill and Gray (1973)

Profelis temmincki (Asian Golden Cat)

527

528

LAURASIATHERIA

Leptailurus serval (Serval)

2n=38

Wurster-Hill and Centerwall (1982)

ORDER CARNIVORA

Octocolobus manul (Pallas’s Cat)

2n=38

Wurster-Hill and Gray (1975)

529

530

LAURASIATHERIA

Callorhinus ursinus (Northern Fur Seal)

2n=36

Arnason (1977)

Arctocephalus pusillus (South African Fur Seal/Southern Fur Seal)

2n=36

Arnason (1977)

ORDER CARNIVORA

Zalophus californianus (California Sea Lion)

2n=36

Arnason (1974b)

2n=36

Arnason (1974b)

Eumetopias jubatus (Northern Sea Lion/Steller Sea Lion)

531

532

LAURASIATHERIA

Odobenus rosmarus (Walrus)

2n=32

Arnason (1977)

2n=34

Arnason (1974b)

Monachus schauinslandi (Hawaiian Monk Seal)

ORDER CARNIVORA

Leptonychotes weddelli (Weddell Seal)

2n=34

Arnason (1974b)

2n=34

Arnason (1974b)

Mirounga angustirostris (Northern Elephant Seal)

533

534

LAURASIATHERIA

Erignathus barbatus (Bearded Seal)

2n=34

Arnason (1974b)

2n=34

Arnason (1974b)

Cystophora cristata (Hooded Seal)

ORDER CARNIVORA

Halichoerus grypus (Gray Seal)

2n=32

Arnason (1970a)

2n=32

Arnason (1974c)

Phoca groenlandica (Harp Seal)

535

536

LAURASIATHERIA

Phoca hispida (Ringed Seal)

2n=32

Arnason (1974c)

Phoca fasciata (Ribbon Seal)

2n=32

Arnason (1977)

ORDER CARNIVORA

Phoca vitulina (Harbor Seal/Common Seal)

2n=32

Nash (unpublished)

# Chromosome identification in brackets is that used by U. Arnason

537

538

LAURASIATHERIA

Ideogram: Phoca vitulina (Harbor Seal/Common Seal)

2n=32

Fröenicke (unpublished)

ORDER PHOLIDOTA

Order Pholidota This order Pholidota contains one family, Manidae, with the single recent genus Manis, distributed in Asia and Africa. There are five subgenera and seven species. At one time pangolins were included with the anteaters, sloths, and armadillos together with the African aardvark in the order Edentata, so named because of the lack of some or all of the teeth (Nowak, 1999). It is now thought, however, that any similarities between the pangolins and other anteating mammals are the results of parallel adaptations to a common way of life and not indicative of actual relationships. Earlier morphological (Rose and Emry, 1993) and biochemical (Sarich, 1993) studies suggested a distant relationships between Pholidota and Carnivora. DNA-sequence-based investigations group Pholidota with Cetartiodactyla, Perissodactyla, Carnivora, Chiroptera, and Eulupotyphla in a superordinal lineage, Laurasiatheria. Within this grouping, Pholidota is placed as sister to the order Carnivora (Murphy et al. 2001a,b, 2002). Pholidota remains one of the least studied orders cytogenetically. As far as is known, most pangolin species have not been karyotyped. The only exception to this is Manis pentadactyla, a species which is widely distributed in South and East Asia. Four diploid numbers (2n = 36, 38, 40, 42) have been reported for M. pentadactyla. The variations in diploid numbers appear to be correlated with differences in the geographical distribution of specimens investigated, 2n = 36 in India (Ray-Chaudhuri et al., 1969; Satya-Prakash and Aswathanarayana, 1972), 2n = 38, 40 in southern China (Feng and Shi, 1983; Quan et al., 1984; Cheng et al., 1991), and 2n = 42 in Japan (Feng and Shi, 1983). A high-quality G-banded karyotype of M. pentadactyla is still not available, but Cheng et al. (1991) suggested that a few tandem fusions could have resulted in reductions in diploid numbers based on variations in the position of the NOR, the relative size of chromosomes, and the centromere index. This atlas presents the G-banded karyotype of Manis javanicus (2n = 38), which has not been reported before. Most recently, we have established a genomewide comparative chromosome map between M. javanicus and human and Martes foina by comparative chromosome painting. The results demonstrate that M. javanicus has a unique karyotype with no cytogenetic signatures that link the order Pholidota to the order Carnivora (Yang personal communication). Fentang Yang

539

540

LAURASIATHERIA

Manis javanica (Javan Pangolin/Malayan Pangolin/ Scaly Anteater)

2n=38

Nie and Yang (unpublished) Contributed by N. W. Nie

ORDER CETARTIODACTYLA

Order Cetartiodactyla Cetacea The traditional order Cetacea, recently included within the newly designated Cetartiodactyla to recognize the polyphyletic evolution of Cetacea and Artiodactyla (Murphy et al., 2001a,b; Springer et al., 2003), includes three suborders, the extinct Arcaeoceti and the recent Odontoceti (toothed whales) and Mysticeti (whalebone or baleen whales). As a result of radical morphological adaptation to the marine (aquatic) environment, it was difficult for a long time to establish the position of Cetacea in the mammalian phylogentic tree and even to settle whether recent cetaceans constituted a monophyletic group. On the basis of biochemical analyses Boyden and Gemeroy (1950) and Goldstone and Smith (1966) had shown a closer relationship between cetaceans and artiodactyls than between cetaceans and any other mammalian order, and comparative karyological studies in the late 1960s and early 1970s unequivocally supported monophyly of recent cetaceans. The nature of the relationship between cetaceans and artiodactyls was resolved in phylogenetic analyses of complete sequences of the mitochondrial cytochrome b gene (Irwin and Arnason, 1994), which placed Cetacea within the order Artiodactyla itself as the sister group of Hippopotamidae (see also Sarich, 1993). This relationship was subsequently supported in analysis of nuclear data (Gatesy et al., 1996; Gatesy, 1997) and statistically established in studies of complete mitochondrial genomes (Ursing and Arnason, 1998). For a more detailed discussion of cetacean relationships the reader should consult Arnason et al. (2004). The first cetacean species of which the karyotype was studied was the Dall porpoise, Phocoenoides dalli, reported by Makino (1948). The analyses were based on direct preparations of male germ cells and the author determined the chromosome number as 2n = 44, including one heteromorphic pair that was selected as the sex chromosomes. The second report on cetacean chromosomes was that of Nowosielski-Slepowron and Peacock (1955), who reported chromosome numbers in the mysticete blue (Balaenoptera musculus) and fin (B. physalus) whales and the odontocete sperm whale (Physter macrocephalus). The analyses of these authors were all made on prophase stages of spermatogonial cells and spermatocytes, and their counts were therefore highly approximate. They estimated the haploid numbers of all three species to 24 but did not find the corresponding diploid number of 48, the highest diploid number counted being 39. The first authors to utilize modern cell culture techniques for chromosome analysis in cetaceans were Atwood and Razavi (1965) and Walen and Madin (1965). Atwood and Razavi (1965) determined the chromosome number of the sperm whale (2n = 42), while Walen and Madin (1965) described the karyotypes of two delphinids, the bottlenosed dolphin (Tursiops truncatus) and the pilot whale (Globicephala macrorhynchus). Both species had 2n = 44 and highly similar karyotypes. Kasuya (1966) reported the chromosome number of the sei whale (Balaenoptera borealis) in direct preparations of spermatogonial cells. The first mysticete karyotype studied with the use of cell culture was that of the fin whale, Balaenoptera physalus (Arnason, 1969). The author presented both male and female karyotypes (2n = 44). A striking feature of the fin whale karyotype was the extreme length of the X chromosome, which measured about 9% of the female haploid set as compared with the standard mammalian length of about 5%. Arnason (1969) advocated monophyly of Odontoceti/ Mysticeti on the basis of the similarities between the karyotype of the mysticete fin whale and the odontocete bottlenosed dolphin and pilot whale. The two major surveys of cetacean cytogenetics (Arnason, 1974a; Duffield,

541

542

LAURASIATHERIA 1977) underlined the slow karyological evolution of Cetacea with essentially the same chromosomal characteristics occurring in different families, such as Balaenopteridae (rorquals), Eschrichtiidae (gray whales), and Delphinidae (dolphins, porpoises). The extraordinary size of the X chromosome of the fin whale compared to that of the dolphin is apparent in this case, but in other respects the similarity between the two karyotypes is striking. The karyological information on marine mammals that had accumulated until 1972 made it possible to propose an explanation to the karyological uniformity characterizing both cetaceans and pinnipeds. The primary causes for this uniformity were formulated by Arnason (1972; see also Arnason, 1982) as being: 1. Low prolificity (a) Late sexual maturity (b) Not more than one offspring per year 2. Good vagility 3. Environment without delimited niches The above factors may be taken as the principal causes of extreme karyotype stability as the chances are remote that individuals having identically rearranged karyotypes will become the founders of new karyotypic subpopulations. The earliest karyological analyses, which for natural reasons were based on unbanded preparations, showed in some instances pronounced size differences within some chromosome pairs. The nature of these observations was not established until the application of chromosome-banding methods. These methods, notably the C-band method, revealed the presence of large C-band positive regions in the cetacean chromosomes, not only in centromeric and terminal positions but also in interstitial regions (Arnason, 1974a). In delphinids the C-band positive regions may amount to 10–15% of the total chromosome length, while in most baleen whales it is 20–25%. The extreme is the fin whale, in which about 30% of the karyotype is C-band positive as the result of the large X chromosome in this species. The nature of the C-band positive regions was later worked out by in situ hybridization experiments. These analyses showed that the common cetacean highly repetitive component (Arnason et al., 1984) occupied essentially all C-bands in the odontocetes (Widegren et al., 1985; Arnason, 1987), while in the mysticetes the common cetacean component primarily occurred in interstitial C-bands (Arnason and Widegren, 1989), with the heavy mysticete satellite (Arnason et al., 1978) occurring in mysticete telomeres and the light mysticete satellite (Arnason et al., 1978) in centromeric regions in all mysticetes other than the right whales (Balaenidae) in which this satellite does not occur (Arnason and Best, 1991). In the following the karyological characteristics of the different families will be discussed. The karyological picture of the mysticetes is more uniform than that of the odontocetes and the account of cetacean cytogenetics will therefore start with the former. The account will focus on conventionally Gand C-banded karyotypes. For information on NORs in cetacean karyotypes the reader might consult Arnason (1981a).

Mysticeti The suborder Mysticeti includes four recent families, Balaenidae, Balaenopteridae, Eschrichtiidae, and Neobalaenidae. Eschrichtiidae (gray whales) was for a long time believed to constitute a basal mysticete lineage, but recent molecular analyses of complete mitochondrial genomes (Arnason et al., 2004) instead identified a basal split between Balaenidae (right whales) and the three

ORDER CETARTIODACTYLA other families among which Neobalaenidae (pygmy right whales) is a sister group to Eschrichtiidae, and Balaenopteridae (rorquals), which share a common branch. Balaenopteridae, Eschrichtiidae, and Neobalaenidae have the common 2n = 44 cetacean karyotype, while Balaenidae, as represented by the bowhead (Balaena mysticetes), has 2n = 42.

Family Balaenopteridae The fin whale was the first mysticete to be cytogenetically analyzed using modern techniques (Arnason, 1969). The general characteristics of the cetacean 2n = 44 karyotype and the presence of four telocentric pairs, three large subtelocentric pairs, and one large submetacentric pair are evident. The X chromosome of the fin whale is exceptionally large and the G-band pattern of the distal part of the long arm of the X chromosome noticeably indistinct. C-bands are prominent in all balaenopterid whales and exceptionally so in the fin whale. In many instances there is also a striking difference in the size of the C-bands in homologous chromosomes. The heterochromatic (Cband positive) nature of the distal part of the long arm of the X chromosome is apparent. The centromeric C-bands of the balaenopterid whales are made up of the so-called light balaenopeterid satellite and the telomeric C-bands of the heavy balaenopterid satellite (Arnason et al., 1978), while interstitial Cbands are made up of the common cetacean highly repetitive component (Arnason and Widegren, 1989). The karyotype of the blue whale, B. musculus, was described by Arnason et al. (1989) and Arnason and Widegren (1989). The X chromosome of the blue whale is of the “standard” placental size. In other respects the karyotype of the blue whale (2n = 44) is highly similar to that of the fin whale. The occurrence of species hybridization between the blue and fin whales has been molecularly demonstrated (Arnason et al., 1991; Spilliaert et al., 1991). It is therefore not excluded that the atypically short X chromosome of the fin whale was a relict of an earlier hybridization between the two species. The karyotype of the minke whale conforms to the common cetacean 2n = 44 karyotype. C-banded karyotypes of the minke whale were described by Arnason et al. (1977; 1978; Arnason, 1974a). A conventionally stained karyotype of the sei whale, B. borealis, was reported by Arnason (1970b). The figure shows also a blowup of the largest pairs of the karyotype. The second pair from the left shows pronounced heteromorphism. The C-band pattern of the sei whale and additional details are described in Arnason (1974a) and Arnason et al. (1978). A Q-banded karyotype of the sei whale was presented in Arnason (1974a). The family Balaenopteridae includes two genera, Balaenoptera and Megaptera. The latter genus has the humpback whale, Megaptera novaeangliae, as its only recent member. The karyotype of this species has not been published before. The karyotype of the humpback whale has the common characteristics of the common 2n = 44 cetacean karyotype. The X chromosome is unusually large. Only the terminal end of the long arm is C-band positive, however. There is apparent heteromorphism in several C-bands.

Family Eschrichtiidae The family Eschrichtiidae contains only one recent species, the gray whale, Eschrichthius robustus. A conventionally stained karyotype of the gray whale was presented by Arnason (1974a) and a Q-banded karyotype by Duffield (1974). The G- and C-banded karyotypes of the gray whale were described by Arnason (1981b). The morphology and banding characteristics of the karyotype of the gray whale conform with those of the balaenopterids.

543

544

LAURASIATHERIA

Family Neobalaenidae The only extant member of the family Neobalaenidae is the pygmy right whale, Caperea marginata. The karyotype of this species has not been published but its conventionally stained karyotype has the chromosome number 2n = 44 and all the morphological characteristics of the mysticete karyotypes discussed above. The presence of the 2n = 44 karyotype in the pygmy right whale is consistent with the phylogenetic position of this species on the same branch as Balaenopteridae and Eschrichtiidae to the exclusion of Balaenidae (Arnason et al., 2004).

Family Balaenidae The family Balaenidae includes two recent species, the bowhead (Balaena mysticetus) and the right whale Balaena (Eubalaena) glacialis. The family Balaenidae is the sister group of remaining mysticetes as demonstrated in various molecular analyses. The light balaenopterid satellite is absent in Balaenidae (Arnason and Best, 1991), and the organization of the heavy satellite in Balaenidae differs from that of the other mysticete families (Adegoke et al., 1993). The conventionally stained and the G-banded karyotypes of the bowhead were described by Jarrell (1979), who suggested that the 2n = 42 karyotype of the bowhead had arisen by fusion of one telomeric pair and one submetacentric pair in the common 2n = 44 mysticete karyotype, thereby giving rise to one large metacentric chromosome and reducing the number of telocentric pairs from four to three. This conclusion was corroborated by in situ hybridization using the heavy mysticete satellite as a probe (Arnason et al., 1982). The study showed that the heavy mysticete satellite, which has terminal chromosome localization in other mysticetes (Arnason et al., 1978), hybridized to an interstitial region in the large metacentric pair identified by Jarrell (1979).

Odontoceti The suborder Odontoceti includes the families Physeteridae, sperm whales, Ziphiidae, beaked whales, Platanistidae, platanistids or Indian river dolphins, and the superfamily Delphinoidea with the families Iniidae, iniid river dolphins, Monodontidae, narwhals and belugas, Phocoenidae, porpoises, and Delphinidae, dolphins. The position of the Yangtze River dolphin, Lipotes vexillifer, has been contentious. Phylogenetic analyses based on complete cytochrome b sequences (Yang et al., 2002) have suggested that it should constitute the monotypic family Lipotidae. That proposal is followed here, but the hypothesis might need substantiation from larger datasets. Delphinoidea and Lipotidae are characterized by the common cetacean 2n = 44 karyotype, while Physeteridae and Ziphiidae have the chromosome number 2n = 42. The ziphiid karyotype resembles the common cetacean karyotype, whereas the relationship between the common cetacean karyotype and the physeterid karyotype has not been worked out. The karyology of Platanistidae is not known. For details of odontocete relationships less Lipotidae, the reader might consult Arnason and Gullberg (1996) and Arnason et al. (2004). C-bands are less conspicuous in the odontocete karyotypes than in the mysticete ones. One reason for this is the virtual absence of both the light (centromeric positions) and the heavy (terminal positions) mysticete DNA satellites in all odontocetes. Molecular analyses based on complete mitochondrial genomes (Arnason et al., 2004) place Physeteridae as the basal sister group of remaining odontocetes. However, for the sake of simplicity the following account will begin

ORDER CETARTIODACTYLA with a discussion of Delphinoidea as its karyotypes conform with the common cetacean karyotype.

Superfamily Delphinoidea The basal delphinoid split is between Iniidae and the three remaining families, Phocoenidae, Monodontidae, and Delphinidae. Among the latter there is a sister group relationship between Phocoenidae and Monodontidae to the exclusion of Delphinidae (Arnason et al., 2004). All cetacean genomes are characterized by the so-called common cetacean highly repetitive component (Arnason et al., 1984). The repeat length of the component is ∼1750 nucleotides (nt) in all cetaceans except Delphinidae, in which the repeat length is ∼1580 nt. This delphinid synapomorphy places the Irrawaddy dolphin, Orcaella brevirostris, solidly among Delphinidae (Gretarsdottir and Arnason, 1992), rather than among Monodontidae, as proposed in some morphological studies. Species belonging to all four delphinoid families have been karyologically described.

Family Iniidae The family Iniidae includes two recent species, Inia geoffrensis, Amazon River dolphin, and Pontoporia blainvillei, La Plata dolphin. The karyotype of the Amazon River dolphin was reported by Duffield (1977). The karyotype has all the characteristics of the 2n = 44 common cetacean karyotype.

Family Monodontidae The family Monodontidae counts two recent species, the narwhal, Monodon monoceros, and the beluga or white whale, Delphinapterus leucas. The conventionally stained and the silver-stained and the G- and C-banded karyotypes of the beluga were described by Jarrell and Arnason (1981). The karyotype has all the characteristics of the 2n = 44 common cetacean karyotype. C-bands are prominent in the beluga and the heteromorphism frequently occurring in homologous C-bands is apparent in pair sm2. This heteromorphism is also evident in the G-banded karyotype. The conventionally stained karyotype of the narwhal was reported by Andrews et al. (1973). Also this karyotype conforms with the common cetacean karyotype.

Family Phocoenidae The karyotype of the harbour porpoise, Phocoena phocoena, was described by Arnason (1974a), who also reported the C-banded karyotype of this species. The karyotype of the harbour porpoise has all the characteristics of the common cetacean 2n = 44 karyotype. Duffield (1977) reported the karyotype of another phocoenoid, Dall’s porpoise, Phocoenoides dalli. The karyotype of this species is the same as that of P. phocoena.

Family Delphinidae Several delphind karyotypes were described by Arnason (1974a) and Duffield (1977). All delphinids have the chromosome number 2n = 44 and, with the exception of the killer whale, the morphology of the karyotypes conforms with the common cetacean 2n = 44 karyotype. In the killer whale the large amount of heterochromatin (C-band positive regions) in the short arms of the four telocentric pairs masks the telocentric nature of these chromosomes. Karyotypes of the common dolphin, Delphinus delphis, Atlantic spinner dolphin, Stenella clymene, bottle-nosed dolphin, Tursiops truncatus, white-beaked

545

546

LAURASIATHERIA dolphin, Lagenorhynchus albirostris, and short-finned pilot whale, Globicephala macrorhynchus, are presented (Arnason, 1974a). The conspicuous heteromorphism, related to the amount of heterochromatin, is evident in pair sm2. The conventionally stained karyotype of a male killer whale, Orcinus orca, is also presented. The four pairs corresponding to the four telocentric pairs in the common 2n = 44 cetacean karyotype have been marked t* in the figure as the telocentric nature of these pairs is masked in the killer whale as the result of accumulation of C-band positive regions in the short arm of these chromosomes. This accumulation is evident in the C-banded karyotype. The C-band heteromorphism occurring in several pairs is also apparent in this figure. Arnason et al. (1980) examined the C-band pattern in six individuals. The pattern was consistent within each individual, whereas the pattern could differ considerably among individuals.

Family Lipotidae As mentioned above the phylogenetic position of Lipotidae may need further examination. The conventionally stained karyotype of the Yangtze River dolphin (beiji), Lipotes vexillifer, was reported by Chen and Chen (1986). The karyotype conforms to the characteristics of the common 2n = 44 cetacean karyotype.

Family Platanistidae No information is currently available on the karyology of this family.

Family Ziphiidae The karyotypes of two ziphiids, Gervais’ beaked whale, Mesoplodon europaeus, and Hubbs’ beaked whale, M. carlhubbsi, were described by Arnason et al. (1977). Both species have 2n = 42 with exceptionally large X chromosomes. The Mesoplodon karyotypes are characterized by the presence of only three pairs of telocentric chromosomes. It is thus probable the Mesoplodon karyotype arose by a fusion of one telocentric pair and one submetacentric pair. The mechanism might thus be analogous to that giving rise to the 2n = 42 balaenid karyotype, although the chromosomes involved might be different. The extraordinary size of the X chromosome in Mesoplodon is due to accumulation of heterochromatin in the distal part of the long arm of this chromosome.

Family Physeteridae Two species belonging to the family Physeteridae, the sperm whale, Physeter macrocephalus, and the pygmy sperm whale, Kogia breviceps, have been karyologically described. Although some similarities exist between the two physeterid karyotypes, their relationship to the common 2n = 44 cetacean karyotype has not been worked out. Common to both karyotypes is the absence of the telocentric chromosomes characterizing other cetacean karyotypes. It may be hypothesized that the m2 pair of the sperm whale karyotype has arisen through fusion of one telocentric pair and the small nuclear organizing pair occurring in the common cetacean 2n = 44 karyotype. Irrespective of the correctness of that hypothesis the C-band pattern shows that the absence of other telocentric chromosomes cannot be explained by accumulation of heterochromatin in the short arms of the original telocentric pairs, as has been the case in the killer whale. Ulfur Arnason

ORDER CETARTIODACTYLA

Lipotes vexillifer (Baiji/Chinese River Dolphin)

2n=44

Minrong et al. (1986)

547

548

LAURASIATHERIA

Delphinapterus leucas (Beluga Whale/White Whale)

2n=44

Jarrell and Arnason (1981) Contributed by U. Arnason

2n=44

Jarrell and Arnason (1981) Contributed by U. Arnason

ORDER CETARTIODACTYLA

Phocoena phocena (Harbor Porpoise/Common Porpoise)

2n=44

Arnason (1980)

2n=44

Arnason (1980)

Stenella clymene (Atlantic Spinner Dolphin)

549

550

LAURASIATHERIA

Stenella attenuata (Pantropical Spotted Dolphin)

2n=44

Stock (1981)

2n=44

Stock (1981)

ORDER CETARTIODACTYLA

Stenella dubia (Spotted Dolphin)

2n=44

Arnason (1974c)

Delphinus delphis (Common Dolphin/Saddlebacked Dolphin)

2n=44

Arnason (1974a)

551

552

LAURASIATHERIA

Lagenorhynchus albirostris (White-Beaked Dolphin)

2n=44

Arnason (1980)

Tursiops truncatus (Bottle-Nosed Dolphin)

2n=44

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Ideogram: Tursiops truncatus (Bottle-Nosed Dolphin)

2n=44

Bielec et al. (1997) Contributed by D. L. Busbee

553

554

LAURASIATHERIA

Orcinus orca (Killer Whale)

2n=44

Arnason et al. (1980)

2n=44

Arnason et al. (1980)

ORDER CETARTIODACTYLA

Globicephala macrorhynchus (Pilot Whale/Blackfish Whale)

2n=44

Arnason (1974a)

2n=42

Arnason et al. (1977)

Mesoplodon carlhubbsi (Hubbs’ Beaked Whale)

555

556

LAURASIATHERIA

Mesoplodon europaeus (Gervais’ Beaked Whale)

2n=42

Arnason et al. (1977)

2n=42

Arnason et al. (1977)

ORDER CETARTIODACTYLA

Kogia breviceps (Pygmy Sperm Whale)

2n=42

Arnason and Benirsche (1973)

557

558

LAURASIATHERIA

Physeter macrocephalus (Sperm Whale)

2n=42

Arnason and Benirsche (1973)

ORDER CETARTIODACTYLA

Eschrichtius robustus (Gray Whale)

2n=44

Arnason (1981a)

Balaena mysticetus (Bowhead Whale/Greenland Right Whale)

2n=42

Jarell (1979) Contributed by U. Arnason

559

560

LAURASIATHERIA

Balaenoptera acutorostrata (Minke Whale)

2n=44

Arnason (1981a)

2n=42

Arnason (1974a)

Balaenoptera physalus (Fin Whale)

ORDER CETARTIODACTYLA

Balaenoptera borealis (Sei Whale)

2n=44

Nash (unpublished)

561

562

LAURASIATHERIA

Balaenoptera musculus (Blue Whale)

2n=44

Arnason and Widegren (1989)

2n=44

Arnason and Widegren (1989)

ORDER CETARTIODACTYLA

Megaptera novaeangliae (Humpback Whale)

2n=44

Arnason (1974a)

563

564

LAURASIATHERIA

Order Cetartiodactyla Artiodactyla Artiodactyla is composed of the even-toed ungulates, and as evidenced by the rich fossil record, this order represents the most significant adaptive radiation among large mammals. The fossil record reveals extensive evolutionary diversity among artiodactyls throughout historical times. The greater than 200 living artiodactyls exhibit a cosmopolitan distribution, absent only from the continents of Australia and Antarctica (Simpson, 1945). Interestingly, molecular evidence has been used to unite the order Cetacea with Artiodactyla. These data suggest that Hippopotamidae and Cetacea are most closely related and should be united with ruminants rather than pigs and peccaries (Nomura et al., 1998; Nomura and Yasue, 1999; for an excellent review of Artiodactyla and its putative phylogenetic relationships with Cetacea, see Lenstra and Bradley, 1999). So, the classical taxonomic treatment of Artiodactyla that follows very well may represent a paraphyletic grouping. Historically, living artiodactyls have been taxonomically subdivided into either 9 (Vaughan, 1986) or 10 (Janis and Scott, 1988) families. Pigs (Suidae, 9 species), peccaries (Tayasuidae, 3 species), and hippopotami (Hippopotamidae, 2 species) are placed in the suborder Suina, whereas camelids (Camelidae, 4 species), chevrotains (Tragulidae, 4 species), deer (Cervidae, 36 species), musk deer (Moschidae, 4 species), giraffids (Giraffidae, 2 species), pronghorn (Antilocapridae, 1 species), and antelope, sheep, goats, bison, and cattle (Bovidae, 124 species) are in the suborder Ruminantia (Vaughan, 1986; animaldiversity.ummz.umich.edu/site/index.html; ultimateungulate.com/ artiodactyla.html). The ruminants are separated into the infraorders Tylopoda (Camelidae) and Pecora (the remaining ruminant families). Pecora consists of the more progressive ruminant families with Tragulidae in many respects being ancestral to the advanced pecoran families (Cervidae/Moschidae, Giraffidae, Antilocapridae, Bovidae). Phylogenetic relationships among the advanced pecorans remain uncertain, and this is reflected in the varied taxonomic placement of these families into superfamilies (Janis and Scott, 1988). Diploid numbers of Suina range from 20 to 38. Suidae (2n = 32–38) has been extensively studied (Benirschke et al., 1985), in part because of significant interest in the economically important domestic pig. A comparative cytogenetic analysis of the three extant species of Tayassuidae (2n = 20–30) revealed a number of conserved chromosomal segments, as well as several segments that could not be paired among species because of pronounced variability in the amount and distribution of heterochromatin (Benirschke et al., 1985; Benirschke and Kumamoto, 1989). The Nile hippopotamus and the pygmy hippopotamus of Hippopotamidae (2n = 36) have essentially identical karyotypes except for Y chromosome differences (Gerneke, 1965; Hsu and Benirschke, 1977). We are not aware of any comprehensive comparative cytogenetic analysis of Suina that has included appropriate out-group comparisons. Diploid numbers of Ruminantia range from 6 to 74. Camelidae (camels, 2n = 74; llama, 2n = 72–74) exhibits the highest diploid numbers. Comparative G- and C-band analysis among South American species and the Bactrian camel revealed what are believed to be identical karyotypes (Bunch et al., 1985). So, it appears that camelids are chromosomally conservative. The family Tragulidae (the chevrotains or mouse deer, 2n = 32; Gallagher et al., 1996) is represented by three extant genera and four species. These small forest dwellers possess a unique mixture of primitive and advanced features and are believed to represent the nearest living relatives of the advanced pecorans (Vaughan, 1986). The advanced pecorans represent approximately 85%

ORDER CETARTIODACTYLA of living artiodactyls with Bovidae (2n = 30–60) and Cervidae/Moschidae (2n = 6–70) comprising about 60 and 20%, respectively. Antilocapridae is monotypic (2n = 58), and Giraffidae is represented by the giraffe (2n = 30) and okapia (2n = 44–46). (See http://www.bionet.nsc.ru/chromosomes/ artiodactyla/artioda2n.htm for an excellent compilation of Artiodactyla karyotypic data and associated references.) Given that the advanced pecorans represent the vast majority of artiodactyls, it is not suprising that they have been extensively studied cytogenetically (Wurster and Benirschke 1967a,b; Wurster and Benirschke, 1968; Evans et al., 1973; Effron et al., 1976; Buckland and Evans, 1978; Fontana and Rubini, 1990; Gallagher and Womack, 1992; Gallagher et al., 1994; Iannuzzi and Di Meo, 1995). These and many other comparative cytogenetic analyses have established extensive monobrachial chromosome conservation among the advanced pecorans and provide insight into the nature of primitive and derivative karyotypes. Despite this, comparative chromosome band analyses have resulted in relatively few character transformations useful in deciphering phylogenetic relationships among families, subfamilies, and tribes. Possibly comparative mapping (Hayes et al., 2003; Iannuzzi et al., 2003; Slate et al., 2002) and molecular cytogenetic (Robinson et al., 1997; Chowdhary et al., 1998; Bielec et al., 1998) approaches coupled with chromosome band analyses will prove of greater utility in understanding genome evolution and phylogenetic relationships among Artiodactyla. Note: Several contributors use Domestic Cattle as the reference species in their numbering system. D.S. Gallagher James Womack

565

566

LAURASIATHERIA

Sus scrofa domestica (Domestic Pig/Hog/Boar)

2n=38

Graphodsky (unpublished)

ORDER CETARTIODACTYLA

Ideogram: Sus scrofa domestica (Domestic Pig/Hog/Boar)

2n=38

Biltueva et al. (2004) Contributed by A. Graphodatsky

Sus scrofa scrofa (European Wild Boar/Hog/Boar)

2n=36

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

567

568

LAURASIATHERIA

Sus barbatus (Bearded Pig/Hog/Boar)

2n=38

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Sus cebifrons (Visayan Warty Pig/Hog/Boar)

2n=34

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Potamochoerus porcus (Red River Hog)

2n=34

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

569

570

LAURASIATHERIA

Potamochoerus larvatus (Southern Bush Pig/African Bush Pig)

2n=34

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Phacochoerus africanus sundervallii (Southern Warthog)

2n=34

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Babyrousa babyrussa celebensis (Sulawesi babirusa)

2n=38

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

571

572

LAURASIATHERIA

Catagonus wagneri (Chacoan Peccary)

2n=20

Benirschke and Kumamoto (1989)

Tayassu pecari (White-Lipped Peccary)

2n=26

Benirschke and Kumamoto (1989)

ORDER CETARTIODACTYLA

Tayassu tajacu (Collared Peccary)

2n=30

Benirschke and Kumamoto (1989)

Hippopotamus amphibius (Hippopotamus)

2n=36

Stanyon (unpublished)

573

574

LAURASIATHERIA

Camelus bactrianus (Bactrian Camel/Two-Humped Camel)

2n=74

Graphodatsky (unpublished)

ORDER CETARTIODACTYLA

Camelus dromedarius (Dromedary Camel/One-Humped Camel)

2n=74

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

575

576

LAURASIATHERIA

Tragulus javanicus (Lesser Malay Chevrotain/Asiatic Mouse Deer)

2n=32

Gallagher et al. (1996)

ORDER CETARTIODACTYLA

Okapia johnstoni (Okapi)

2n=45–46

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Giraffa camelopardalis tippelskirchi (Masai Giraffe)

2n=30

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)* * In memory of Arlene T. Kumamoto for her many contributions to cytogenetics of endangered species.

577

578

LAURASIATHERIA

Moschus moschiferus (Siberian Musk Deer)

2n=58

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Elaphodus cephalophus (Tufted Deer)

2n=46–48

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Muntiacus muntjak vaginalis (North Indian Muntjac/Barking Deer)

2n=6(F) 2n=7(M)

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

579

580

LAURASIATHERIA

Muntiacus feae (Fea’s Muntjac/Barking Deer)

2n=14 XY

Fu and Yang (unpublished) Contributed by F. Yang

Muntiacus reevesi (Chinese muntjac/Barking Deer)

2n=46

Yang (unpublished)

ORDER CETARTIODACTYLA

Ideogram: Muntiacus reevesi (Chinese Muntjac/Barking Deer)

2n=46

Yang et al. (1995)

581

582

LAURASIATHERIA

Muntiacus gongshanensis (Gongshan Muntjac/Barking Deer)

2n=8 XX

Fu and Yang (unpublished)

Muntiacus crinifrons (Black Muntjac/Barking Deer)

Contributed by F. Yang

Female

Male

2n=8 XX 2n=9 XY

Fu and Yang (unpublished) Contributed by F. Yang

ORDER CETARTIODACTYLA

Dama mesopotamica (dama) (Persian Fallow Deer)

2n=68

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

583

584

LAURASIATHERIA

Axis kuhlii (Kuhl’s Deer/Axis Deer)

2n=68

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Axis calamianensis (Calamian Deer/Axis Deer)

2n=68

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

585

586

LAURASIATHERIA

Cervus unicolor malaccensis (Malayan Sambar Deer/Wapiti)

2n=56

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Cervus nippon hortulorum (Dybowski’s Sika)

2n=66

Graphodatsky (unpublished)

587

588

LAURASIATHERIA

Cervus nippon mandarinus (Mandarin Sika)

2n=66

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Cervus albirostris (White-Lipped Deer/Thorold’s Deer)

2n=66

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

589

590

LAURASIATHERIA

Cervus elaphus bactrianus (Bactrian Wapiti/Red Deer)

2n=68

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Cervus elaphus sibiricus (Red Deer/Wapiti)

2n=68

Graphodatsky (unpublished)

591

592

LAURASIATHERIA

Alces alces (Moose/Elk)

2n=68

Graphodatsky (unpublished)

ORDER CETARTIODACTYLA

Rangifer tarandus (Siberian Reindeer/Caribou)

2n=70

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

593

594

LAURASIATHERIA

Capreolus pygargus (Roe Deer)

2n=70 + B’s

Graphodatsky (unpublished)

ORDER CETARTIODACTYLA

Antilocapra americana (Pronghorn)

2n=58

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

595

596

LAURASIATHERIA

Tragelaphus angasii (Lowland Nyala)

2n=55(M)

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Tragelaphus strepsiceros (Greater Kudu)

2n=32

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

597

598

LAURASIATHERIA

Tragelaphus imberbis (Lesser Kudu)

2n=38

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Tragelaphus eurycerus (Bongo)

2n=34(F) 2n=33(M)

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Taurotragus oryx pattersonianus (Patterson’s Eland)

2n=32(F) 2n=31(M)

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

599

600

LAURASIATHERIA

Taurotragus derbianus (Giant Eland)

2n=32(F) 2n=31(M)

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Boselaphus tragocamelus (Nilgai)

2n=46

Gallagher et al. (1998)

601

602

LAURASIATHERIA

Bubalus bubalis (Asian Water Buffalo)

2n=50

Iannuzzi (1994)

ORDER CETARTIODACTYLA

Ideogram: Bubalus bubalis (Asian Water Buffalo)

2n=50

Iannuzzi (1994)

603

604

LAURASIATHERIA

Bubalus depressicornis (Lowland Anoa)

2n=48

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Syncerus caffer (Forest Buffalo/African Buffalo)

2n=54–56

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

605

606

LAURASIATHERIA

Bos taurus (Domestic Cattle)

2n=60

Graphodatsky and Biltueva (unpublished)

ORDER CETARTIODACTYLA

Ideogram: Bos taurus (Domestic Cattle)

2n=60

Cribiu et al. (2001) Contributed by D. S. Gallagher

Positive G (left) and R (right) bands Negative G (left) and R (right) bands Centromeric regions

607

608

LAURASIATHERIA

Bos javanicus (Javan Banteng/Bali Cattle)

2n=60

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Bos frontalis (gaurus) (Gaur/Seladang)

2n=58

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

609

610

LAURASIATHERIA

Bos grunniens (Yak/Oxen)

2n=60

Graphodatsky and Biltueva (unpublished)

ORDER CETARTIODACTYLA

Bison bonasus (European Bison/Wisent)

2n=60

Graphodatsky (unpublished)

611

612

LAURASIATHERIA

Cephalophus rufilatus (Red-Flanked Duiker)

2n=60

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Cephalophus niger (Black Duiker)

2n=60

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

613

614

LAURASIATHERIA

Cephalophus sylvicultor (Yellow-Backed Duiker)

2n=60

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Cephalophus dorsalis (Bay Duiker)

2n=60

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

615

616

LAURASIATHERIA

Cephalophus monticola (Blue Duiker)

2n=60

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Kobus ellipsiprymnus defassa (Defassa Waterbuck)

2n=53–54

Kingswood et al. (1998a) Contributed by O. A. Ryder

Kobus ellipsiprymnus ellipsiprymnus (Ellipsen Waterbuck)

2n=50–52

Kingswood et al. (1998a) Contributed by O. A. Ryder

617

618

LAURASIATHERIA

Kobus megaceros (Nile Lechwe)

2n=52

Kingswood et al. (2000) Contributed by M. L. Houck

ORDER CETARTIODACTYLA

Kobus leche (Red Lechwe)

2n=48

Kingswood et al. (2000) Contributed by M. L. Houck

619

620

LAURASIATHERIA

Kobus kob thomasi (Uganda Kob)

2n=50

Kingswood et al. (2000) Contributed by M. L. Houck

ORDER CETARTIODACTYLA

Pelea capreolus (Rhebok)

2n=56

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

621

622

LAURASIATHERIA

Hippotragus equinus (Roan Antelope)

2n=60

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Hippotragus niger (Sable Antelope)

2n=60

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

623

624

LAURASIATHERIA

Oryx dammah (Scimitar-Horned Oryx)

2n=56–58

Kumamoto et al. (1999) Contributed by O.A. Ryder

ORDER CETARTIODACTYLA

Oryx leucoryx (Arabian Oryx)

2n=57–58

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

625

626

LAURASIATHERIA

Oryx gazella beisa (Beisa Oryx)

2n=58

Kumamoto et al. (1999) Contributed by O. A. Ryder

ORDER CETARTIODACTYLA

Oryx gazella callotis (Fringe-Eared Oryx)

2n=58

Kumamoto et al. (1999) Contributed by O. A. Ryder

627

628

LAURASIATHERIA

Oryx gazella gazella (Gemsbok)

2n=56

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Addax nasomaculatus (Addax)

2n=58

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

629

630

LAURASIATHERIA

Damaliscus hunteri (Hunter’s Hartebeest)

2n=44

Kumamoto et al. (1996) Contributed by M. L. Houck

ORDER CETARTIODACTYLA

Damaliscus pygargus phillipsi (Blesbok/Sassabie)

2n=38

Kumamoto et al. (1996) Contributed by M. L. Houck

631

632

LAURASIATHERIA

Damaliscus lunatus topi (Topi)

2n=36

Kumamoto et al. (1996) Contributed by M. L. Houck

ORDER CETARTIODACTYLA

Alcelaphus buselaphus jacksoni (Jackson’s Hartebeest)

2n=40

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

633

634

LAURASIATHERIA

Connochaetes gnou (White-Tailed Gnu/Black Wildebeest)

2n=58

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Connochaetes taurinus albojubatus (Brindled Gnu)

2n=58

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Oreotragus oreotragus stevensoni (Klipspringer)

2n=60

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

635

636

LAURASIATHERIA

Raphicerus campestris (Steenbok)

2n=30

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Neotragus moschatus (Suni Antelope/Dwarf Antelope)

2n=52–56

Kingswood et al. (1998b) Contributed by S. J. Charter

ORDER CETARTIODACTYLA

Madoqua guentheri (Guenther’s Dik-Dik)

2n=48–50

Kingswood and Kumamoto (1996)

637

638

LAURASIATHERIA

Madoqua kirkii (Kirk’s Dik-Dik)

2n=46–48

Kingswood and Kumamoto (1997)

ORDER CETARTIODACTYLA

Antilope cervicapra (Blackbuck)

2n=30–33

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

639

640

LAURASIATHERIA

Aepyceros melampus petersi (Black-Faced Impala)

2n=58–60

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Gazella saudiya (Saudi Gazelle)

2n=46–53

Kumamoto et al. (1995)

Gazella bennettii (Chinkara/Gazelle/Indian Gazelle)

Contributed by M. L. Houck

2n=49–52

Kumamoto et al. (1995) Contributed by M. L. Houck

641

642

LAURASIATHERIA

Gazella gazella (Mountain Gazelle)

2n=34(F) 2n=35(M)

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Gazella spekei (Speke’s Gazelle)

2n=32(F) 2n=33(M)

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Gazella cuvieri (Cuvier’s Gazelle)

2n=32(F) 2n=33(M)

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Gazella rufifrons laevipes (Red-fronted Gazelle)

2n=58

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

643

644

LAURASIATHERIA

Gazella subgutturosa subgutturosa (Persian Goitered Gazelle)

2n=32(F) 2n=31(M)

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Gazella subgutturosa marica (Sand Gazelle)

2n=30–32F 2n=31–33M

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Gazella thomsonii (Thomson’s Gazelle)

2n=58

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Gazella granti roosevelti (Roosevelt’s Gazelle)

2n=30(F) 2n=31(M)

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

645

646

LAURASIATHERIA

Gazella dama ruficollis (Addra Gazelle)

2n=38–40

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Gazella dama mhorr (Mhorr Gazelle)

2n=38–40

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Gazella soemmerringii (Soemmerring’s Gazelle)

2n=34–39

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

647

648

LAURASIATHERIA

Antidorcas marsupialis (Springbok/Springbuck)

2n=56

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

Oreamnos americanus (Rocky Mountain Goat)

2n=42

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Capricornis crispus (Serow)

2n=50

Biltueva Graphodatsky and Sen-ichi (unpublished) Contributed by A. Graphodatsky

649

650

LAURASIATHERIA

Naemorhedus goral (Goral)

2n=56

Biltueva Graphodatsky and Sen-ichi (unpublished) Contributed by A. Graphodatsky

ORDER CETARTIODACTYLA

Budorcas taxicolor (Takin)

2n=52

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

651

652

LAURASIATHERIA

Ovibos moschatus (Muskox)

2n=48

Biltueva and Graphodatsky Contributed by A. Graphodatsky

ORDER CETARTIODACTYLA

Capra hircus (Domestic Goat)

2n=60

Biltueva and Graphodatsky Contributed by A. Graphodatsky

653

654

LAURASIATHERIA

Capra cylindricornis (East Caucasian Tur/Goat)

2n=60

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Capra pyrenaica hispanica (Spanish Ibex/Goat)

2n=60

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

655

656

LAURASIATHERIA

Pseudois nayaur szechuanensis (Chinese Bharal/Blue Sheep)

2n=56

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER CETARTIODACTYLA

Ovis orientalis musimon (European Mouflon/Sheep)

2n=54

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

657

658

LAURASIATHERIA

Ovis aries (Domestic Sheep)

2n=54

Cribiu et al. (2001) Contributed by D. S. Gallagher

ORDER CETARTIODACTYLA

Ideogram: Ovis aries (Domestic Sheep)

2n=54

Cribiu et al. (2001) Contributed by D. S. Gallagher

659

660

LAURASIATHERIA

Ovis canadensis (Rocky Mountain Bighorn Sheep)

2n=54

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER PERISSODACTYLA

Order Perissodactyla The order Perissodactyla, as the odd-toed hoofed mammals are known, is characterized by the relative enlargement of digit III on each extremity. Other morphological characteristics also distinguish this group, which is a sister taxon to the even-toed ungulates, Artiodactyla. The order contains three living groups: horses, tapirs, and rhinoceroses. Equidae is known from the earliest Eocene, while tapirs and rhinoceroses differentiated in the late Eocene from a chalicothere ancestor. All 16 extant species of Perissodactyla are threatened or endangered except Equus quagga and one subspecies of Ceratotherium simum. As a result, access to specimens for study, especially from individuals living in the wild, is extremely limited. Despite this difficulty, G-banded karyotypes have been described for all but one species in this order, the Javan rhinoceros (Rhinoceros sondaicus). Diploid numbers range from 32 to 84, and intraspecific numerical polymorphisms have been observed in a surprisingly large number of perissodactyls. Robertsonian fission/fusion events are prevalent in both Equidae and Rhinocerotidae, often involving the same two acrocentric pairs across species. De novo fission has been documented in three equid taxa: plains zebra (Whitehouse et al., 1984), domestic donkey (TrommershausenBowling and Millon, 1988), and Somali wild ass (Houck et al., 1998). Recently, cross-species chromosome painting has identified autosomal segments conserved between zebras and the African rhinoceroses (Trifonov et al., 2003). Similar studies comparing conserved chromosomal segments among tapirs, horses, and rhinoceroses are underway. Ultimately they will fill in the picture of the evolution of the perissodactyl karyotype.

Family Equidae Groves and Ryder (2000) list seven extant species of Equidae. The subgenus Equus includes a single species, Equus ferus, which includes the Przewalski’s (Bowling et al., 2004) and domestic horses. The Przewalski’s horse (E. f. przewalskii) has 66 chromosomes (Benirschke et al., 1965), whereas domestic horses (E. f. caballus) have 2n = 64 chromosomes. The subgenus Asinus consists of the hemionus group (Asian wild asses), which includes Equus hemionus and Equus kiang, each of which have identified subspecies. Diploid numbers of 54 and 55 have been described in E. h. kulan and 55–56 in E. h. onager; both forms share the same Robertsonian polymorphism (Ryder, 1978; Ryder and Chemnick, 1990). Analysis of equid mtDNA suggests paraphyly and implicates a recent separation, casting doubt on their validity as separate subspecies (Oakenfull et al., 2000). Diploid numbers of 51 and 52 have been observed in E. kiang; the fission/fusion polymorphism resulting in the numerical variation leads to the expectation that 2n = 50 karyotypes might also exist, although they have not yet been described in the limited population studies reported to date. The asinus group (African asses) consists of a single species, Equus africanus, which includes two wild subspecies and the domestic donkey. Diploid numbers in E. a. somaliensis, the Somali wild ass, range from 62 to 64 due to a centric fission/fusion event (Houck et al., 1998) involving chromosomes homologous to those in E. kiang, E. h. kulan, E. h. onager and E. quagga (burchelli) boehmi (Myka et al., 2003b). The X chromosomes of E. africanus differ from those of all other equids by a pericentric inversion (Houck et al., 1998). The subgenus Hippotigris includes the extant zebras. Equus zebra, which includes two subspecies, has the lowest chromosome number among the equids with a diploid number of 32. The common or plains

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LAURASIATHERIA zebra, Equus quagga, has four commonly recognized subspecies. The first reports of the plains zebras, E. quagga (Hansen, 1975a; Ryder et al., 1978), identified the diploid number to be 2n = 44. However, Whitehouse et al. (1984) later identified a fusion polymorphism resulting in an individual with 2n = 45. The Grevy’s zebra, E. grevyi, has 2n = 46. Comparative G-banding studies suggested homologies between elements in the equid karyotype (Ryder et al., 1978), but it was 25 years later that the hypothesized rearrangements were examined by comparative mapping and Zoo-FISH. By FISH mapping horse-derived gene markers, Myka et al. (2003a) confirmed that the karyotypes of E. f. caballus and E. f. przewalskii differed by a single fission/fusion event involving domestic horse chromosome 5 and Przewalski’s horse chromosomes 23 and 24. At the same time, Yang et al. (2003b) arrived at the same conclusions using cross-species chromosome painting. Studies demonstrating the presence of interstitial telomere sites in E. q. boehmi and E. z. hartmannae have provided further evidence of chromosome fusion events contributing to equid chromosome evolution (Lear, 2001; Santani et al., 2002). The comparative cytogenetics studies have suggested features of the phylogeny of equids, such as the close relationship between E. f. caballus and E. f. przewalskii, the monophyly of zebras, and the close relationship between E. quagga and E. grevyi. The chromosomal rearrangements of E. z. hartmannae were too complex to elucidate with Gbanding studies but have now been undertaken using chromosome painting (Richard et al., 2001; Yang et al., 2003b). Studies of molecular evolution of mitochondrial DNA RFLPs (Restriction Fragment Length Polymorphism) (George and Ryder, 1986) and, later, mitochondrial DNA sequences (Oakenfull et al., 2000) suggest that E. z. hartmannae is closely related to the other two extant zebra species and that zebras are monophyletic. Consequently, the complex rearrangements that differentiate the karyotype of E. zebra from E. grevyi and E. quagga must have taken place relatively recently. Equids are notable for their rapid rate of chromosomal evolution (Bush et al., 1977; Wichman et al., 1991) and for their ability to produce viable interspecies hybrids from parents with differentiated karyotypes (Benirschke and Malouf, 1967; Ryder et al., 1978; Gray, 1972). Hybrids between Przewalski’s horse and domestic horses are fertile, although most other equine hybrids that have been evaluated are sterile (Ryder et al., 1978). Studies of mitochondrial DNA of Przewalski’s and domestic horses show a high degree of similarity, with Przewalski’s horse haplotypes nested within the variation observed among breeds of domestic horses (Ishida et al., 1995; Oakenfull and Ryder, 1998; Vila et al., 2001). Studies of Y chromosome sequence variation in equids have established that the Y chromosomes of Przewalski’s and domestic horses do not share a recent common ancestor (Wallner et al., 2003). Introgression between domestic horses and Przewalski’s horses has been identified several times in the studbook population of the endangered Przewalski’s horse; nonetheless, analysis of nuclear genetic variation indicates that these forms represent identifiably distinct gene pools (Bowling et al., 2004). No discussion of equine hybrids and their cytogenetics would be complete without mention of the mule, notorious for its sterility, but with persistent reports of an occasional fertile female mule (Benirschke and Ryder, 1985). Recently, detailed differences between the karyotypes of the domestic horse and the domestic donkey have been elucidated through chromosome painting by Raudsepp and Chowdhary (1999). In 1985 a case of a fertile female mule was presented that documented the pedigree and qualified the offspring to the mule and her donkey mate (Ryder et al., 1985). The chromosomes of the offspring were those of a mule, and nuclear markers also provided evidence that only maternal (horse) alleles were transmitted by the female mule. As yet,

ORDER PERISSODACTYLA there is no good explanation of how unprecedented segregation of parental genetic material might have occurred.

Family Rhinocerotidae Four of the five extant species that comprise this family have been karyotyped, with only the elusive, critically endangered Javan rhinoceros (Rhinoceros sondaicus) remaining unstudied. Numerous small acrocentric chromosomes presented challenges in determining consistent diploid numbers for this family during the infancy of endangered species cytogenetics (Hansen, 1975b; Heinichen, 1967, 1970; Hungerford et al., 1967; Wurster-Hill and Benirschke, 1968; Hsu and Benirschke, 1973). Later G-band studies identified the diploid numbers as 2n = 82 for the Sumatran (Dicerorhinus sumatrensis) and Indian (Rhinoceros unicornis) rhinoceroses and 2n = 84 for the black rhinoceros (Diceros bicornis) (Houck et al., 1994, 1995). However, centric fusion of two acrocentric chromosomes was noted in three related Ceratotherium simum cottoni individuals, resulting in a diploid number of 2n = 81 (Houck et al., 1994). The G-banding pattern of the rhinoceros X chromosome is similar to that of the standard type submetacentric X found in other mammalian species, including humans, for all species in this group. The X chromosome of D. sumatrensis has a block of heterochromatin at the distal portion of the long arm, cytogenetically similar to the X chromosome of the mountain tapir, Tapirus pinchaque. The most parsimonious explanation for this finding would be that the interstitial heterochromatic block is an independently derived character. However, the relationship between heterochromatic blocks, repetitive DNA sequences, and chromosomal dynamics is far from clearly depicted at this time and synapomorphic events cannot be strictly precluded (Pevzner and Tesler, 2003). Size polymorphisms involving the largely heterochromatic short arms of many autosomes are prevalent among Rhinocerotidae species, making it difficult to assign a static number of acrocentric versus submetacentric chromosomes per species, yet also serving as a measurable difference distinguishing subspecies in some instances (i.e., Diceras bicornis minor vs. D. b. michaeli). As in Equidae, Robertsonian translocations are prevalent and have been identified in all rhinoceros species studied thus far, with the exception of R. unicornis (Houck et al., 1994, and unpublished data). To date, only the heterozygous form of the fission/fusion mechanism has been documented in rhinoceroses, while both heterozygous and homozygous conditions have been noted in some equid species (i.e., Equus africanus somaliensis and E. hemionus). Future FISH studies will be helpful in determining if the elements involved in Rhinocerotidae are the same as those identified in equids.

Family Tapiridae This family is represented by a single genus, Tapirus, and four species that primarily occur in Central and South America with the exception of the Malayan tapir (T. indicus), which is found in Southeast Asia. Although nondifferentially stained and C-banded karyotypes for the South American or lowland tapir (T. terrestris) were published in 1975 by Hsu and Benirschke, G-banded karyotypes of the four tapir species were not available until recently (Houck et al., 2000). The diploid numbers are 2n = 52 for T. indicus, 2n = 76 in T. pinchaque (mountain tapir), and 2n = 80 in both T. terrestris and T. bairdii (Baird’s tapir). Although the latter two species have the same diploid number, their karyotypes are distinguishable even without banding because of the single biarmed autosomal pair in T. terrestris compared to eight pairs in the T. bairdii karyotype. The number of autosome arms (NAA) documented in the four tapir species varies from 80 to 94, suggesting that chromosome rearrangements other than

663

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LAURASIATHERIA centric fissions/fusions were involved in the karyotypic evolution of this family. There are at least 13 conserved autosomes between the karyotypes of T. bairdii and T. terrestris, while at least 15 are conserved between T. bairdii and T. pinchaque and at least 17 between T. terrestris and T. pinchaque (Houck et al., 2000). There appear to be fewer homologies between the Malayan tapir and the three species inhabiting Central and South America. As in Rhinocerotidae, the G-banding pattern of the X chromosome is similar to that of the human X, and the T. pinchaque X chromosome exhibits a block of heterochromatin at the distal portion of the long arm, similar to that observed in the Sumatran rhinoceros (Houck et al., 2000). Marlys L. Houck Oliver A. Ryder

ORDER PERISSODACTYLA

Equus asinus (Donkey)

2n=62

Alaoui et al. (2004) Contributed by M. Ponsa

Equus asinus (africanus) somaliensis (Somali Wild Ass)

2n=62–64

Houck et al. (1998)

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LAURASIATHERIA

Equus hemionus onager (Onager)

2n=55–56

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER PERISSODACTYLA

Equus hemionus kulan (Kulan)

2n=54–55

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

667

668

LAURASIATHERIA

Equus przewalskii (Przewalski’s Wild Horse)

2n=66

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER PERISSODACTYLA

Equus caballus (Domestic Horse)

2n=64

Stanyon (unpublished)

669

670

LAURASIATHERIA

Ideogram: Equus caballus (Domestic Horse)

2n=64

Bowling et al. (1997) Contributed by F. A. Ponce de Leon

ORDER PERISSODACTYLA

Equus burchelli (Burchell’s Zebra)

2n=44

Graphodatsky (unpublished)

671

672

LAURASIATHERIA

Equus quagga boehmi (Grant’s Zebra/Quagga)

2n=44–45

Center for Reproduction of Endangered Species (CRES) Zoological Society of San Diego (unpublished)

ORDER PERISSODACTYLA

Tapirus terrestris (Lowland Tapir)

2n=80

Houck et al. (2000)

673

674

LAURASIATHERIA

Tapirus pinchaque (Mountain Tapir)

2n=76

Houck et al. (2000)

ORDER PERISSODACTYLA

Tapirus bairdii (Baird’s Tapir)

2n=80

Houck et al. (2000)

675

676

LAURASIATHERIA

Tapirus indicus (Malayan Tapir)

2n=52

Houck et al. (2000)

ORDER PERISSODACTYLA

Rhinoceros unicornis (Indian Rhinoceros)

2n=82

Houck et al. (1994)

677

678

LAURASIATHERIA

Ceratotherium simum simum (Southern White Rhinoceros/SquareLipped Rhinoceros)

2n=82

Houck et al. (1994)

ORDER PERISSODACTYLA

Diceros bicornis (Black Rhinoceros)

2n=84

Houck et al. (1995)

679

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INDEX Note: Chromosome pictures and ideograms are denoted by bold page numbers. A Aardvark, 80 Abrocoma bennetti, 334 Abrothrix longipilis, 196 Abrothrix sanborni, 196 Acinonyx jubatus, 523 Aconaemys fuscus, 333 Aconaemys porteri, 333 Aconaemys sagei, 333 Acrobates pygmaeus, 36, 62 Acrobatidae, 9, 36 Addax, 629 Addax nasomaculatus, 629 Aepyceros melampus petersi, 640 Afrosoricidia, 63 Afrotheria, 63 Agouti paca, 174 Agouti taczanowskii, 174 Ailuropoda melanoleuca, 447, 479, 480 Ailurus fulgens, 447, 481 Akodon cursor, 174 Akodon longipillis, 196 Akodon olivaceus, 196 Akodon sanborni, 196 Akodon simulator, 195 Akodon toba, 195 Alcelaphus buselaphus jacksoni, 633 Alces alces, 592 Alionycteris paucidentata, 387 Allenopithecus nigroviridis, 150 Allocricetulus curtatus, 208 Allocricetulus eversmanni, 207 Alopex lagopus, 455, 456 Alouatta belzebul, 120, 121 Alouatta caraya, 122 Alouatta sara, 119 Alouatta seniculus, 118 Alticola argentatus, 238

Alticola macrotis, 239 Amblysomus hottentotus, 63, 67 Amblysomus iris, 66 Amblysomus julianae, 66 Amblysomus robustus, 63 Amblysomus septentrionalis, 63 Ametrida centurio, 425 Amphinectomys savamis, 194 Anoa: lowland, 604 Anoura caudifera, 411 Anoura cultrata, 411 Anteater: giant, 85; lesser, 86; scaly, 540; silky, 87 Antechinomys laniger, 26 Antechinus flavipes, 22 Antelope: dwarf, 636; Roan, 622; sable, 623; Suni, 636 Antidorcas marsupialis, 648 Antilocapra americana, 595 Antilocapridae, 564 Antilope cervicapra, 639 Aotus azarae, 127 Aotus nancymaae, 127 Apes, great, 100 Apes, lesser, 100 Aplodontia rufa, 173 Apodemus peninsulae, 175, 279 Apodemus sylvaticus, 280 Archboldomys luzonensis, 285 Arctictis binturong, 503 Arctocephalus pusillus, 530 Ardops nichollsi, 423 Ariteus flavescens, 425 Armadillo: hairy, 88; La Plata three-banded, 92; longnosed, 92, 93; naked-tailed, 91; six-banded, 89 Artibeus jamaicensis, 423 Artibeus lituratus, 424 Artiodactyla, 564 Arvicanthis ansorgei, 304 Arvicanthis niloticus-Cytotype 1a, 302

Atlas of Mammalian Chromosomes, Edited by Stephen J. O’Brien, Joan C. Menninger, William G. Nash Copyright © 2006 John Wiley & Sons, Inc.

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Arvicanthis niloticus-Cytotype 1b, 303 Arvicanthis rufinus, 305 Arvicanthis sp., 306 Arvicola terrestris, 241 Ass: Somali wild, 665 Atelocynus microtis, 463 Atilax paludinosus, 508 Atopogale cubanus, 357 Auliscomys boliviensis, 198 Auliscomys micropus, 198 Auliscomys sublimis, 198 Axis calamianensis, 585 Axis kuhlii, 584 Aye-aye, 115 B Baboon: hamadryas, 153 Babyrousa babyrussa celebensis, 571 Badger: ferret, 498; honey, 497; old world, 498 Baiji, 547 Balaena glacialis, 544 Balaena mysticetus, 543, 544, 559 Balaenidae, 544 Balaenoptera acutorostrata, 560 Balaenoptera borealis, 561 Balaenoptera musculus, 541, 562 Balaenoptera physalus, 541, 560 Balaenopteridae, 543 Bandicoot: rabbit-eared, 32; long-nosed, 34; shortnosed, 33 Banteng: Javan, 608 Bassaricyon sp., 484 Bassariscus astutus, 447, 481 Bat: African long-tongued fruit, 388; Allen’s big-eared, 440; American epauleted, 416, 417; American longeared, 439; Asian false vampire, 391, 392; Australian big-eared, 442; bamboo, 435, 436; big brown, 437, 438; big-eared, 402; big-eyed, 420, 421, 422; Blandford’s, 435; bonneted, 443, 444; broad-faced, 442, 443; brown flower, 413; bulldog, 399; chestnut long-nosed, 410; chocolate, 435; club-footed, 435, 436; common vampire, 400; Cuban flower, 414; dark long-tongued, 412; dawn, 388; Dormer’s, 434; dwarf epauleted, 386; dwarf little fruit, 415; epauleted, 385; falcate-winged, 425; Fischer’s little fruit, 416; fisherman’s, 399; frog-eating, 405; funnel-eared, 427; Geoffroy’s long-nosed, 411; Godman’s long-nosed, 413; Gray’s spear-nosed, 408; groove-lipped, 435; hairy-legged vampire, 401; hairy-tailed, 440; hammerheaded fruit, 385; Hart’s little fruit, 423; hog-nosed, 412; horseshoe, 392, 393; house, 437, 438, 439; Indiana, 428; Jamaican fig-eating, 425; lappet-eared, 440; Leach’s single-leaf, 409; leaf-chinned, 399; leaflipped, 397, 398; lesser broad-nosed, 436; Linnaeus’s false vampire, 405; little big-eared, 402, 403, 404; little brown, 427, 428; little collared fruit, 383; little white-shouldered, 425; little yellow-faced, 422; lobelipped, 435; long-tailed, 388; long-tongued, 409; lump-nosed, 439; mastiff, 443, 444; Mexican longnosed, 412; mouse-tailed, 388; moustached, 397, 398;

naked-backed, 397, 398; neotropical fruit, 423, 424; new world free-tailed, 443; New Zealand short-tailed, 426; old world leaf-nosed, 394, 395, 396; Peter’s spear-nosed, 406; Peter’s woolly false vampire, 406; pinto, 441; red, 440; red fruit, 426; round-eared, 408; Rousette fruit, 381, 382; Saussure’s long-nosed, 410; serotine, 437, 438; short-tailed leaf-nosed, 414, 415; spear-nosed, 407; spectral vampire, 405; spotted, 441; tent-building, 417; tomb, 390, 391; tree, 423; tubenosed insectivora, 441; Underwood’s long-tongued, 412; velvety free-tailed, 444; visored, 426; wattled, 435; white, 422; white-lined, 418, 420, 421, 422; white-winged vampire, 401; yellow, 438, 439; yellow shoulder, 416, 417; yellow-eared, 419 Bathyergus janetta, 319 Bdeogale sp., 507 Bear: American black, 474; ant, 80; Asiatic black, 473; brown, 475; polar, 476; sloth, 478; spectacled, 471, 472; sun, 477 Beaver, 182, 183 Bharal: Chinese, 656 Bilby, 32 Binturong, 503 Bison: European, 611 Bison bonasus, 611 Blackbuck, 639 Blanfordimys afghanus, 245 Blanfordimys bucharicus, 246 Blarina carolinensis, 370 Blarina carolinensis penninsulae, 371 Blarina hylophaga, 371 Blesbok, 631 Boar: European wild, 567 Bongo, 599 Bonobo, 170 Bos frontalis, 609 Bos gaurus, 609 Bos grunniens, 610 Bos javanicus, 608 Bos taurus, 606, 607 Boselaphus tragocamelus, 601 Bovidae, 564 Brachylagus idahoensis, 349 Bradypodidae, 81 Brushtail: common, 37 Bubalus bubalis, 602, 603 Bubalus depressicornis, 604 Budorcas taxicolor, 651 Buffalo: African, 605; Asian water, 602, 603; forest, 605 Bunolagus monticularis, 347 Burramyidae, 9, 36 Burramys parvus, 36 Bushbaby, 108 C Cabassous centralis, 91 Cabassous tatouay, 91 Cacajao calvus rubicundus, 124 Cacomistle, 481 Caenolestidae, 19

INDEX Calcochloris obtusirostris, 63 Callicebus cupreus, 125 Callicebus pallescens, 126 Callicebus personatus nigrifrous, 126 Callimico goeldii, 102, 132 Callithrix argentata, 134 Callithrix chrysoleuca, 137 Callithrix emiliae, 135 Callithrix humeralifer, 136 Callithrix jacchus, 134 Callithrix mauesi, 137 Callorhinus ursinus, 530 Calomyscus bailwardi, 202 Calomyscus mystax, 200 Calomyscus urartensis, 201 Caluromyidae, 9, 10 Caluromys lanatus, 10, 17 Caluromys philander, 10 Camel: Bactrian, 574; dromedary, 575; one-humped, 575; two-humped, 574 Camelidae, 564 Camelus bactrianus, 574 Camelus dromedarius, 575 Campagnol, 253 Campagnol des Champs, 269 Canidae, 446 Canis familiaris, 450, 467, 468 Canis latrans, 465 Canis lupus, 466 Caperea marginata, 544 Capra cylindricornis, 654 Capra hircus, 653 Capra pyrenaica hispanica, 655 Capreolus pygargus, 594 Capricornis crispus, 649 Capromys pilorides, 174 Capuchin: weeper, 131 Capybara, 325 Caracal, 527 Caracal caracal, 527 Caribou, 593 Carnivora, 445 Carollia brevicauda, 414 Carollia castanea, 414 Carollia perspicillata, 415 Casiragua, 335, 336, 337, 338 Castor canadensis, 173, 183 Castor fiber, 173, 182 Cat: Asian golden, 527; Asian leopard, 525; black-footed, 517; domestic, 514, 515; fishing, 526; Geoffroy’s, 513; jungle, 516; little spotted, 511; Pallas’s, 529; Pampas, 512; sand, 516; tiger, 511 Catagonus wagneri, 572 Cattle: Bali, 608; domestic, 606, 607 Cavia cobaya, 324 Cavia porcellus, 324 Cavies, 324 Cebuella pygmaea, 138 Cebus albifrons albifrons, 129 Cebus apella, 130

Cebus capucinus, 128 Cebus olivaceus, 131 Celaenomys silaceus, 285 Cephalophus dorsalis, 615 Cephalophus monticola, 616 Cephalophus niger, 613 Cephalophus rufilatus, 612 Cephalophus sylvicultor, 614 Ceratotherium simum, 661 Ceratotherium simum simum, 678 Cercartetus concinnus, 36, 58 Cercartetus lepidus, 36 Cercartetus nanus, 36 Cercocebus galeritus, 151 Cercocebus torquatus, 152 Cercopithecus aethiops, 141, 142 Cercopithecus ascanius schmidti, 144 Cercopithecus cephus, 146 Cercopithecus diana, 147 Cercopithecus l’hoesti, 148 Cercopithecus mona campbell, 145 Cercopithecus nictitans, 143 Cercopithecus petaurista, 145 Cercopithecus pogonias, 147 Cerdocyon thous, 458, 459 Cerocebus torquatus, 152 Cervidae, 564 Cervus albirostris, 589 Cervus elaphus bactrianus, 590 Cervus elaphus sibiricus, 591 Cervus nippon hortulorum, 587 Cervus nippon mandarinus, 588 Cervus unicolor malaccensis, 586 Cetacea, 541 Cetartiodactyla, 541, 564 Chaetophractus villosus, 88 Chalinolobus morio, 435 Cheetah, 523 Chevrotain: lesser Malay, 576 Chimpanzee: common, 168, 169; pygmy, 170 Chinchilla laniger, 174 Chinchillone, 334 Chinkara, 641 Chionomys gud, 244 Chionomys nivalis, 242 Chionomys roberti, 243 Chipmunk, 177 Chiroderma doriae, 421 Chiroderma improvisum, 422 Chiroderma villosum, 420 Chironectes minimus, 10 Chiroptera, 378 Chlorotalpa duthieae, 65 Chlorotalpa sclateri, 65 Chlorococebus aethiops, 141 Choeroniscus godmani, 413 Choeroniscus intermedius, 413 Choeronycteris mexicana, 412 Choloepus didactylus, 83 Choloepus hoffmanni, 84

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INDEX

Chroeomys olivaceus, 196 Chrotopterus auritus, 406 Chrysochloris asiatica, 63 Chroeomys olivaceus, 196 Chrysocyon brachyurus, 469 Civet: African palm, 500; banded palm, 505; lesser Oriental, 502; Malagasy, 505; masked palm, 504; palm, 503 Clethrionomys californicus, 234 Clethrionomys centralis, 236 Clethrionomys gapperi, 234 Clethrionomys glareolus, 235 Clethrionomys rufocanus, 231 Clethrionomys rutilus, 233 Clethrionomys sikotanensis, 232 Colobus guereza, 102, 159 Colugo, 98, 99 Conie, 345, 346 Connochaetes gnou, 634 Connochaetes taurinus albojubatus, 634 Coro-coro, 340 Coruro, 332 Corynorhinus townsendii, 439 Cottontail, 350; Eastern, 351; mountain, 352; New England, 351 Cougar, 522 Coyote, 465 Coypu, 341 Cricetulus barabensis, 176, 210 Cricetulus griseus, 212 Cricetulus longicaudatus, 214 Cricetulus migratorius, 209 Cricetulus obscurus, 213 Cricetulus pseudogriseus, 211 Cricetulus triton, 219 Cricetus cricetus, 206 Crocidura dsinezumi, 376 Crocidura horsfieldii watasei, 373 Crocidura pergrisea, 374 Crocidura sibirica, 374 Crocidura suaveolens, 373 Crocidura yankariensis, 359 Crocuta crocuta, 449, 510 Crunomys suncoides, 284 Cryptomys hottentotus damarensis, 321 Cryptomys mechowi, 174, 320 Cryptoprocta ferox, 508 Ctenodactylus gundi, 312 Ctenodactylus vali, 313 Ctenomys fulvus, 174, 328 Ctenomys magellanicus magellanicus, 327 Ctenomys maulinus brunneus, 326 Ctenomys maulinus maulinus, 327 Ctenomys opimus, 327 Ctenomys rionegrensis, 325 Ctenomys robustus, 326 Ctenomys talarum, 326 Cu Lan, 106, 107 Cyclopes didactylus, 87 Cynocephalidae, 98

Cynocephalus volans, 98, 99 Cystophora cristata, 534 D Dactylomys dactylinus, 340 Dama dama, 583 Dama mesopotamica, 583 Damaliscus hunteri, 630 Damaliscus lunatus topi, 632 Damaliscus pygargus phillipsi, 631 Dassie: rock, 79 Dasymys incomtus, 300 Dasymys rufulus, 301 Dasypodidae, 82 Dasypus hybridus, 93 Dasypus novemcinctus, 92 Dasypus septemcinctus, 93 Dasyuridae, 9, 21 Dasyuroides byrnei, 27 Dasyuromorphia, 9, 21 Dasyurus hallucatus, 28 Dasyurus maculatus, 30 Dasyurus viverrinus, 29 Daubentonia madagascariensis, 115 Deer: Asiatic mouse, 576; axis, 584, 585; barking, 579, 580, 581, 582; Calamian, 585; Kuhl’s, 584; Malayan sambar, 586; Persian fallow, 583; red, 590, 591; roe, 594; Siberian musk, 578; Thorold’s, 589; tufted, 579; white-lipped, 589 Degus, 328, 329 Delphinapterus leucas, 545, 548 Delphinidae, 545 Delphinoidea, 545 Delphinus delphis, 545, 551 Dendrohyrax arboreus, 78 Dermoptera, 98 Desmana moschata, 358 Desmodus rotundus, 400 Diaemus youngi, 401 Diceras bicornis michaeli, 663 Diceras bicornis minor, 663 Dicerorhinus sumatrensis, 663 Diceros bicornis, 663, 679 Dicrostonyx torquatus, 175, 274 Didelphidae, 9, 10 Didelphimorphia, 9, 10 Didelphis marsupialis, 18 Didelphis virginiana, 18 Dik-dik: Guenther’s, 637; Kirk’s, 638 Dinomys branickii, 323 Diphylla ecaudata, 401 Dipodomys merriami, 187 Diprotodontia, 9, 36 Distoechurus pennatus, 36 Dog: bush, 464; Chinese raccoon, 462; domestic, 467, 468; Japanese raccoon, 460, 461; round-eared, 463; small-eared, 463 Dolphin: Atlantic spinner, 549; bottle-nosed, 552, 553; Chinese river, 547; common, 551; pantropical spotted, 550; saddlebacked, 551; spotted, 551; white-beaked, 552

INDEX Donkey, 665 Dormouse: common, 314; edible, 315; fat, 315; forest, 316; garden, 317; mouselike, 318 Dromiciops australis, 20 Dryomys nitedula, 316 Duiker: bay, 615; black, 613; blue, 616; red-flanked, 612; yellow-backed, 614 E Echidna: long-beaked, 4, 7; short-beaked, 3, 6 Echinops telfairi, 69 Echymipera kalabu, 31 Ectophylla alba, 422 Eira barbara, 495 Eland: greater, 600; Patterson’s, 599 Elaphodus cephalophus, 579 Elephant: African forest, 74; African savannah, 74, 77; Asian/Indian, 74, 76 Elephantidae, 74 Elephantulus bracyrhynchus, 64 Elephantulus edwardii, 64 Elephantulus intufi, 64 Elephantulus myurus, 64 Elephantulus rupestris, 64 Elephas maximus, 74, 76 Eligmodontia puerulus, 197 Eligmodontia sp., 197 Eliomys quercinus, 317 Elk, 592 Ellobius fuscocapillus, 173, 278 Ellobius lutescens, 279 Ellobius talpinus, 174, 175, 277 Ellobius tancrei, 175 Enchisthenes hartii, 423 Eonycteris robusta, 388 Eothenomys regulus, 237 Epomops franqueti, 385 Eptesicus bottae, 437 Eptesicus fuscus, 438 Equidae, 661 Equus africanus, 661 Equus africanus somaliensis, 661, 663, 665 Equus asinus, 665 Equus asinus somaliensis, 665 Equus burchelli, 671 Equus caballus, 669, 670 Equus ferus, 661 Equus ferus caballus, 661, 662 Equus ferus przewalskii, 662 Equus greyvi, 662 Equus hemionus, 661 Equus hemionus kulan, 661, 667 Equus hemionus onager, 661, 666 Equus kiang, 661 Equus przewalskii, 668 Equus quagga, 661, 662 Equus quagga boehmi, 662, 672 Equus zebra, 661 Equus zebra hartmannae, 662 Erethizon dorsatum, 174

705

Erignathus barbatus, 534 Erinaceidae, 358 Erinaceus amurensis, 362 Erinaceus concolor, 361 Erinaceus europaeus, 360 Ermine, 484 Erophylla sezekorni, 413 Erythrocebus patas, 140 Eschrichitius robustus, 543, 559 Eschrichtiidae, 543 Euarchontoglires, 95 Eubalaena glacialis, 544 Euderma maculatum, 441 Eulemur fulvus sanfordi, 111 Eulemur macaco flavifrons, 110 Eulipotyphla, 357 Eumetopias jubatus, 531 Eumops glaucinus, 444 Eumops perotis, 443 Euneomys chinchilloides petersoni, 199 Euneomys mordax, 199 Euphractus sexcinctus flavimanus, 89 F Falsistrellus tasmaniensis, 433 Felidae, 450 Felis catus, 514, 515 Felis chaus, 516 Felis margarita, 516 Felis nigripes, 517 Fennecus zerda, 454 Ferret: domesticated, 490 Fossa, 508 Fossa fossana, 449, 505 Fox: Arctic, 455, 456; bat-eared, 470; Corsac, 453; crabeating, 458, 459; Fennec, 454; flying, 384; grey, 457; Indian flying, 384; kit, 453; long-haired flying, 387; long-haired tailless flying, 387; red, 451, 452; Rodriguez flying, 383 G Galago, 108; brown, 109 Galago moholi, 108 Galea musteloides, 174 Galemys pyrenaicus, 358 Galeopterus variegatus, 98 Galictis vittata, 495 Galidia elegans, 506 Gaur, 609 Gazella bennetti, 641 Gazella cuvieri, 643 Gazella dama mhorr, 646 Gazella dama ruficollis, 646 Gazella gazella, 642 Gazella granti roosevelti, 645 Gazella rufifrons laevipes, 643 Gazella saudiya, 641 Gazella soemmerringii, 647 Gazella spekei, 642 Gazella subgutturosa marica, 644

706

INDEX

Gazella subgutturosa subgutturosa, 644 Gazella thomsonii, 645 Gazelle: Addra, 646; Cuvier’s, 643; Indian, 641; Mhorr, 646; mountain, 642; Persian goitered, 644; redfronted, 643; Roosevelt’s, 645; sand, 644; Saudi, 641; Soemmerring’s, 647; Speke’s, 642; Thomson’s, 645 Gemsbok, 628 Genet: large spotted, 501; small spotted, 501 Genetta genetta, 501 Genetta tigrina, 449, 501 Geocapromys brownii, 174 Geomys attwateri, 186 Georychus capensis, 319 Gerbil: Congo, 228; large naked-soled, 222; northern pygmy, 225; Petter’s, 226; rocky, 225; Senegal, 226, 227; small naked-soled, 226, 227, 228, 229; Tranier’s, 227 Gerbillus nigeriae, 175 Gerbillus rupicola, 225 Gibbon: black-crested, 160, 161; concolor, 160, 161; silvery, 163; white-banded, 162 Giraffa camelopardalis tippelskirchi, 577 Giraffe: Masai, 577 Giraffidae, 564 Glider: feather-tailed, 36; squirrel, 37; sugar, 37; yellowbellied, 37 Glironiidae, 11 Globicephala macrorhynchus, 541, 546, 555 Glossophaga soricina, 409 Gnu: brindled, 634; white-tailed, 634 Goat: domestic, 653; east Caucasian, 654; rocky mountain, 648; Spanish, 655 Gopher: pocket, 186; western pocket, 184 Goral, 650 Gorilla: lowland, 166, 167 Gorilla gorilla, 166, 167 Grammomys gazellae, 173 Grammomys macmillani, 173 Grisón: Allamand’s, 495; greater, 495 Guenon, 147; greater spot-nosed, 143; L’Hoest, 148; lesser spot-nosed, 145; moustached, 146; Schmidt’s spot-nosed, 144 Guinea pig, 324 Gulo gulo, 497 Gundi: atlas, 312; Sahara, 313 Gymnobelideus leadbeateri, 37 Gymnure: Philippine, 360 H Halichoerus grypus, 535 Hamster: Afghan mouselike, 200; black-bellied, 206; Brandt’s, 216, 217; Campbell’s, 204; Chinese, 212; ciscaucasian, 220; Dzhungarian, 203; Eversmann’s, 207; Gobi, 213; gray dwarf, 209; lesser long-tailed, 214; long-tailed, 215; Mongolian, 208; mouselike, 202; Romanian, 221; small desert, 205; striped dwarf, 210; Syrian golden, 218, 219; Transbaikal dwarf, 211; Urartsk mouselike, 201 Hare: black-tailed jack, 353; mouse, 345, 346 Hartebeest: Hunter’s, 630; Jackson’s, 633

Hedgehog: Amur, 362; Brandt’s, 363; Daurian, 365; desert, 363; eastern Eurasian, 361; Eurasian, 362; long-eared desert, 364; small Madagascar, 69; western Eurasian, 360 Helgale pavula, 449 Heliophobius argenteocinereus, 174 Helogale parvula, 507 Hemibelideus lemuroides, 38, 60 Hemicentetes nigriceps, 64, 68 Hemiechinus auritus, 364 Hemiechinus dauricus, 365 Hemigalus derbyanus, 505 Herpailurus yagouaroundi, 524 Herpallura yagouaroundi, 450 Herpestes javanicus, 506 Herpestidae, 449 Hesperoptenus blandfordi, 435 Heterocephalus glaber, 174 Heterohyrax brucei, 78 Hippopotamus, 573 Hippopotamus amphibius, 573 Hipposideros ater, 394 Hipposideros cineraceus, 395 Hipposideros fulvus, 394 Hipposideros fulvus pallidus, 395 Hipposideros lankadiva, 395 Hipposideros pomona, 394 Hipposideros speoris, 396 Hippotragus equinus, 622 Hippotragus niger, 623 Hog: Red River, 569 Homo sapiens, 171, 172 Horse: domestic, 669, 670; Przewalski’s wild, 668 Human, 171, 172 Hyaenidae, 449 Hydrochaeris hydrochaeris, 174, 325 Hyena: brown, 509; spotted, 510 Hylobates concolor, 101, 160, 161 Hylobates hoolock, 101 Hylobates lar, 162 Hylobates moloch, 163 Hylobates syndactylus, 101, 160 Hylonycteris underwoodi, 412 Hypsignathus monstrosus, 385 Hypsiprymnodon moschatus, 38 Hypsugo crassulus, 431 Hypsugo eisentrauti, 434 Hyracoidea, 78 Hyrax, 79 I Ibex: Spanish, 655 Icthyomys pittieri, 173 Ictonyx striatus, 496 Idionycteris phyllotis, 440 Impala: black-faced, 640 Iniidae, 545 Isodon obesulus, 33 Isothrix sinnamariensis, 339

INDEX J Jaguarundi, 524 K Kagwang, 98, 99 Kangaroo: eastern gray, 55; grey, 37; musky rat, 38; red, 37, 57; western gray, 56 Kikuya, 159 Kinkajou, 482 Klipspringer, 635 Koala, 37, 40 Kob: Uganda, 620 Kobus ellipsiprymnus defassa, 617 Kobus ellipsiprymnus ellipsiprymnus, 617 Kobus kob thomasi, 620 Kobus leche, 619 Kobus megaceros, 618 Kogia breviceps, 546, 557 Kowari, 27 Kudu: greater, 597; lesser, 598 Kulan, 667 Kultarr, 26 L Lagenorhynchus albirostris, 546, 552 Lagomorpha, 342 Lagorchestes conspicillatus, 37 Lagostomus maximus, 174, 322 Lagothrix lagotricha, 118 Lagurus lagurus, 272 Langur: Douc, 157 Lasionycteris noctivagans, 379 Lasiorhinus latifrons, 39, 42 Lasiurus borealis, 440 Laurasiatheria, 357 Lechwe: Nile, 618; red, 619 Lemming: bog, 275; collared, 274; steppe, 272; varying, 274 Lemmus sibiricus, 275 Lemur: black, 110; flying, 98, 99; mouse, 110; northern sportive, 113; red-tailed, 112; red-tailed sportive, 112; Sanford’s, 111; white-footed sportive, 112 Leontopithecus chrysopygus, 139 Leontopithecus rosalia, 139 Leopard: Bornean clouded, 520; mainland clouded, 521 Leopardus pardalis, 511 Leopardus tigrinus, 511 Leopoldamys edwardsi, 287 Lepilemur leucopus, 112 Lepilemur ruficaudatus, 112 Lepilemur septentrionalis, 113 Leporidae, 342 Leptailurus serval, 528 Leptonychotes weddelli, 533 Lepus californicus, 353 Lepus saxatilis, 354 Lichonycteris obscura, 412 Linsang: banded, 502; Oriental, 502 Lion, 518; mountain, 522 Lionycteris spurrelli, 410

707

Lipotes vexillifer, 544, 546, 547 Lipotidae, 546 Lissonycteris angolensis, 382 Lonchophylla robusta, 410 Lonchophylla thomasi, 410 Lonchotrix emiliae, 339 Lophuromys brevicaudus, 298 Lophuromys chrysopus, 299 Lophuromys melanonyx, 297 Loris: pygmy slow, 106; slender, 105; slow, 107 Loris tardigradus, 105 Loxodonta africana, 74, 77 Loxodonta cyclotis, 74 Loxodontomys micropus, 198 Lutra lutra, 500 Lynchailurus colocolo, 512 Lynx: Eurasian, 524 Lynx lynx, 524 M Macaca fascicularis, 152 Macaca fuscata, 154, 155 Macaca mulatta, 153 Macaque: crab-eating, 152; Japanese, 154, 155 Macropodidae, 9, 36 Macropus eugenii, 37, 52, 53 Macropus fuliginosus, 37, 56 Macropus giganteus, 37, 55 Macropus parma, 51 Macropus parryi, 54 Macropus robustus, 37, 57 Macropus rufogriseus, 51 Macropus rufus, 37, 57 Macroscelidea, 63 Macroscelides proboscideus, 64, 70 Macrotis lagotis, 32 Macrotus waterhousii, 402 Macrotys waterhousii, 380 Madoqua guentheri, 637 Madoqua kirkii, 638 Manatee: Florida, 72, 73 Mandrill, 156 Mandrillus sphinx, 156 Mangabey: collared, 152; red-capped, 152; Tana River, 151 Manidae, 539 Manis javanica, 540 Manis javanicus, 539 Manis pentadactyla, 539 Maresomy boliviensis, 198 Marmosa sp., 12 Marmoset: common, 134; Emilia’s, 134; Goeldi’s, 132; gold and white, 137; Maues, 137; pygmy, 138; Santarem, 136; silvery, 134 Marmosidae, 9, 11 Marmosops incanus, 12 Marsupalia, 9 Marsupial “mouse”: fat-tailed, 25; flat-skulled, 23; yellow footed, 22 Marsupial mole, 35

708

INDEX

Marten: beach, 491; Japanese, 494; pine, 493; yellowthroated, 494 Martes flavigula, 494 Martes foina, 491, 539 Martes martes, 493 Martes melampus, 494 Martes zibellina, 493 Massoutiera, 311 Massoutiera mzabi, 311 Mastomys awashensis, 290 Mastomys erythroleucus-Cytotype 3, 289 Mastomys huberti, 291 Mastomys kollmanspergeri, 292 Mastomys natalensis, 288 Megaderma lyra, 392 Megaderma spasma, 391 Megaloglossus woermanni, 388 Megalonychidae, 81 Megaptera novaeangliae, 543, 563 Meles meles, 498 Mellivora capensis, 497 Melogale sp., 498 Menetes berdmorei, 173, 176 Mesechinus dauricus, 365 Mesocricetus auratus, 218, 219 Mesocricetus brandti, 216, 217 Mesocricetus newtoni, 221 Mesocricetus raddei, 220 Mesophylla macconnelli, 422 Mesoplodon carlhubbsi, 546, 555 Mesoplodon europaeus, 546, 556 Metachirus nudicaudatus, 14 Meteoro de Prado, 269 Micoureus demerarae, 13 Microbiotheria, 9, 20 Microbiotheriidae, 9, 20 Microcebus murinus, 110 Microgale dobsoni, 68 Micronycteris brachyotis, 404 Micronycteris hirsuta, 404 Micronycteris megalotis, 402 Micronycteris minuta, 379, 403 Micronycteris nicefori, 404 Micronycteris schmidtorum, 403 Micropotamogale dobsoni, 64 Micropteropus pusillus, 386 Microtus agrestis, 269 Microtus arvalis arvalis, 258 Microtus breweri, 270 Microtus californicus, 271 Microtus daghestanicus, 249 Microtus evoronensis, 267 Microtus fortis, 263 Microtus gregalis, 268 Microtus guentheri arm, 256 Microtus guentheri philistinus, 257 Microtus hyperboreus, 264 Microtus juldaschi carruthersi, 247 Microtus kirgisorum, 261

Microtus longicaudus, 254 Microtus majori, 250 Microtus maximowiczii, 266 Microtus mexicanus, 272 Microtus mongolicus, 264 Microtus montebelli, 262 Microtus mujanensis, 266 Microtus nasarovi, 251 Microtus ochrogaster, 253 Microtus oeconomus, 262 Microtus oregoni, 173, 271 Microtus pennsylvanicus, 269 Microtus pinetorum, 253 Microtus richardsoni, 254 Microtus rossiaemeridionalis, 259 Microtus sachalinensis, 265 Microtus schelkovnikovi, 252 Microtus socialis, 255 Microtus socialis schiddlovskii, 256 Microtus sp. nova from iran, 273 Microtus subterraneus, 248 Microtus townsendii, 270 Microtus transcaspicus, 260 Mimon crenulatum, 408 Mink: American, 488, 489; European, 487 Miopithecus talapoin, 149 Mirounga angustirostris, 533 Mole: African golden, 65; Cape golden, 63; hairy-tailed, 377; long-clawed, 276; old world, 376, 377; Siberian, 377; South African golden, 66, 67; southern, 278; transcaucasian, 279 Mole-rat: African, 223; big-headed, 223; east African, 224; Mediterranean blind, 222 Molossops abrasus, 443 Molossops temminckii, 442 Molossus ater, 444 Molossus molossus, 444 Molossus rufus, 444 Monachus schauinslandi, 532 Mongoose: African marsh, 508; black-legged, 507; dwarf, 507; Java, 506; Malagasy ring-tailed, 506; water, 508 Monito del Monte, 20 Monkey: African green, 141, 142; Allen’s swamp, 150; Atlantis titi, 126; Azara’s night, 127; Azara’s whitefaced, 128; black and white colobus, 159; black howler, 122; Bolivian red howler, 119; coppery titi, 125; Diana, 147; dusty leaf, 156; L’Hoest, 148; Mona, 145; Nancy Ma’s night, 127; Patas, 140; red howler, 118; red-handed howler, 120, 121; red-tailed, 144; rhesus, 153; silver leaf, 158; squirrel, 131; tufted capuchin, 130; white-coated titi, 126; white-fronted capuchin, 129; white-nosed, 143; wooly, 118 Monodelphis domestica, 15, 16 Monodontidae, 545 Monophyllus redmani, 409 Monotremata, 1 Moose, 592 Mormoops blainvillii, 399

INDEX Moschidae, 564 Moschus moschiferus, 578 Mouflon: European, 657 Mouse: Andean big-eared, 198; birch, 187; Bolivian bigeared, 198; brush, 190; brush-furred, 297, 298, 299; cactus, 188; Chako grass, 195; Dalton’s, 293; deer, 188, 189, 190, 191; grey-bellied grass, 195; hazel, 314; highland desert, 197; house, 307, 308; Korean wood, 279; long-haired grass, 196; long-tailed field, 280; Macedonian, 309; Mahomet, 309; old world wood, 279, 280; olive grass, 196; Patagonian chincilla, 199; piñon, 191; Sanborn’s grass, 196; southern bigeared, 198; white-footed, 189, 190, 191; white-footed deer, 188 Muntiacus crinifrons, 582 Muntiacus feae, 580 Muntiacus gongshanensis, 582 Muntiacus muntjak vaginalis, 579 Muntiacus reevesi, 580, 581 Muntjac: black, 582; Chinese, 580, 581; Fea’s, 580; Gongshan, 582; North Indian, 579 Murina cyclotis, 441 Mus goundae, 173 Mus macedonicus, 309 Mus mahomet, 309 Mus musculus, 174, 176, 307, 308 Mus platythrix, 176 Muscardinus avellanarius, 314 Muskox, 652 Muskrat: round-tailed, 274 Mustela altaica, 486 Mustela erminea, 484 Mustela eversmanni, 491 Mustela lutreola, 487 Mustela nivalis, 485 Mustela putorius furo, 490 Mustela putorius putorius, 489 Mustela sibirica, 487 Mustela sibirica itatsi, 488 Mustela vison, 488, 489 Mustela visons, 448 Mustelidae, 447 Myocastor coypus, 174, 341 Myomimus personatus, 174, 318 Myomys daltoni, 293 Myonycteris torquata, 383 Myopus schisticolor, 176 Myotis keenii, 427 Myotis mystacina, 428 Myotis nigricans, 428, 429 Myotis sodalis, 428 Myotis thysanodes, 427 Myoxus glis, 315 Myrmecobiidae, 21 Myrmecobius fasciatus, 21 Myrmecophaga tridactyla, 85 Myrmecophagidae, 82 Mystacina tuberculata, 426 Mysticeti, 542

N Naemorhedus goral, 650 Nandinia binotata, 448, 500 Nandiniidae, 448 Nannospalax ehrenbergi, 222 Natalus major, 427 Neobalaenidae, 544 Neofelis nebulosa, 521 Neofelis nebulosa diardi, 520 Neofiber alleni, 274 Neomys fodiens, 372 Neotoma albigula, 188 Neotragus moschatus, 636 Nesokia indica, 286 Nilgai, 601 Ningaui sp., 24 Ningauis, 24 Noctilio albiventris, 399 Notoryctemorphia, 35 Notoryctes typhlops, 35 Notoryctidae, 35 Nutria, 341 Nyala: lowland, 596 Nyctereutes procyonoides procyonoides, 462 Nyctereutes procyonoides viverrinus, 460, 461 Nycticebus coucang, 106, 107 Nycticebus pygmaeus, 106 Nyctinomops laticaudatus, 443 Nyctophilus gouldi, 442 O Ocelot, 511 Ochotona alpina, 343, 344 Ochotona collaris, 344 Ochotona cuniculus, 344 Ochotona curzoinae, 344 Ochotona daurica, 343, 344 Ochotona hyperborea, 344, 346 Ochotona pallasi, 343, 344 Ochotona princeps, 343, 344, 345 Ochotona pusilla, 343, 344 Ochotona rujescens, 343, 344 Ochotona rutila, 343 Ochotonidae, 343 Octocolobus manul, 529 Octodon degus, 329 Octodon lunatus, 328 Odobenidae, 448 Odobenis rosmarus, 448 Odobenus rosmarus, 532 Odontoceti, 544 Oilgoryzomys flavescens, 193 Okapi, 577 Okapia johnstoni, 577 Oligoryzomys flavescens, 193 Oligoryzomys microtis, 193, 193 Olingo, 484 Onager, 666 Oncifelis geoffroyi, 513

709

710

INDEX

Opossum: black four-eyed, 17; brown four-eyed, 14; Chilean “shrew,” 19; common, 18; elegant fat-tailed, 13; mouse, 12, 13; short-tailed, 15, 16; South American mouse, 12; Virginia, 18; wooly, 10, 17 Orangutan: Sumatran, 164, 165 Orcaella brevisrostris, 545 Orcinus orca, 546, 554 Oreamnos americanus, 648 Oregailarus jacobita, 450 Oreotragus oreotragus stevensoni, 635 Ornithorhynchus anatinus, 3, 8 Orycteropodidae, 80 Orycteropus afer, 80 Oryctolagus cuniculus, 343, 354, 355 Oryx: Arabian, 625; Beisa, 626; fringe-eared, 627; scimitar-horned, 624 Oryx dammah, 624 Oryx gazella beisa, 626 Oryx gazella callotis, 627 Oryx gazella gazella, 628 Oryx leucoryx, 625 Oryzomys megacephalus, 192 Oryzomys nitidus, 192 Oryzomys yunganus, 190 Otariidae, 448 Otocyon megalotis, 470 Otolemur crassicaudatus, 109 Otopteropus cartilagonodus, 387 Otter: European river, 500 Ovibos moschatus, 652 Ovis aries, 658, 659 Ovis canadensis, 660 Ovis orientalis musimon, 657 Oxen, 610 P Pacarana, 323 Pademelon: red-bellied, 45; red-necked, 46 Paguma larvata, 449, 504 Pan paniscus, 170 Pan troglodytes, 168, 169 Panda: giant, 479, 480; lesser, 481; red, 481 Pangolin: Javan, 540; Malayan, 540 Panthera leo, 518 Panthera tigris altaica, 519 Papio hamadryas, 153 Paradoxurus hermaphroditus, 503 Paraechinus hypomelas, 363 Parahyaena brunnea, 449, 509 Parascalops breweri, 377 Paucituberculata, 19 Peccary: Chacoan, 572; collared, 573; white-lipped, 572 Pectinator: Speke’s, 311 Pectinator spekei, 311 Pedetes capensis, 310 Pelea capreolus, 621 Peramelemorphia, 9, 31 Perameles nasuta, 34 Peramelidae, 9, 31 Perissodactyla, 661

Perognathus hispidus, 173 Peromyscus boylii, 190 Peromyscus eremicus, 188 Peromyscus maniculatus, 189 Peromyscus truei, 191 Peroryctidae, 31 Petauridae, 9, 37 Petaurista albiventer, 181 Petaurus australis, 37 Petaurus breviceps, 37 Petaurus norfolcensis, 37, 61 Petrodomus tetradactylus, 64 Petrogale brachyotis, 49 Petrogale concinna, 49 Petrogale lateralis lateralis, 47 Petrogale penicillata, 48 Petrogale xanthopus celeris, 46 Petrogale xanthopus xanthopus, 47 Petropseudes dahli, 60 Phacochoerus africanus sundervallii, 571 Phalangeridae, 9, 37 Phascolarctidae, 37 Phascolarctos cinereus, 37, 40 Phenacomys intermedius, 240 Philander opossum, 17 Phloeomys cumingi, 284 Phoca fasciata, 536 Phoca groenlandica, 535 Phoca hispida, 536 Phoca vitulina, 448, 537, 538 Phocidae, 448 Phocoena phocoena, 545, 549 Phocoenidae, 545 Phocoenoides dalli, 541, 545 Phodopus roborovskii, 205 Phodopus sungorus campbelli, 204 Phodopus sungorus sungorus, 203 Pholidota, 539 Phylloderma stenops, 406 Phyllonycteris poeyi, 414 Phyllops haitiensis, 425 Phyllostomus discolor, 407 Phyllostomus hastatus, 407 Physeter macrocephalus, 541, 546, 558 Physeteridae, 546 Pichi, 90 Pig: African bush, 570; bearded, 568; domestic, 566, 567; southern bush, 570; Visayan warty, 568 Pika: American, 345; northern, 346 Pipistrelle, 429, 430, 431, 432, 433, 434 Pipistrellus ceylonicus, 430 Pipistrellus coromandra, 430 Pipistrellus crassulus, 431 Pipistrellus eisentrauti, 434 Pipistrellus javanicus, 430 Pipistrellus kuhli, 431 Pipistrellus mimus, 433 Pipistrellus pipistrellus, 429 Pipistrellus sagittula, 432 Pipistrellus stenopterus, 432

INDEX Pipistrellus tasmaniensis, 433 Pipistrellus vulturnus, 432 Pithecia pithecia, 123 Planigale maculata, 23 Platanistidae, 546 Platypus, 3, 8 Platyrrhinus vittatus, 418 Platyrrhinus vittatus lineatus, 418 Podogymnura truei, 360 Polecat: marbled, 492; steppe, 491; striped, 496; wild, 489 Pongo pygmaeus abelii, 164, 165 Pontoporia blainvillei, 545 Porpoise: common, 549; harbor, 549 Possum: common brush-tailed, 43; common ring-tailed, 38, 59; Daintree River ringtail, 38; feather-tailed, 36; Herbert River ringtail, 38; honey, 61; Leadbeater’s, 37; lemuroid ring-tailed, 38, 60; little eastern, 36; little pygmy, 36; mountain pygmy, 36; pygmy gliding, 62; rock ring-tailed, 60; western pygmy, 36, 58 Potamochoerus larvatus, 570 Potamochoerus porcus, 569 Potoroidae, 9, 38 Potoroo: long-footed, 38; long-nosed, 44 Potorous gilberti, 38 Potorous longipes, 38 Potorous tridactylus, 38, 44 Potos flavus, 447, 482 Primates, 100 Prionailurus bengalensis, 525 Prionailurus viverrinus, 526 Prionodon linsang, 502 Proboscidea, 74 Procavia capensis, 78, 79 Procyon lotor, 447, 482, 483 Procyonidae, 447 Proechimys brevicauda-Cytotype 1, 335 Proechimys simonsi, 337 Proechimys sp. 1, 338 Proechimys sp. 2, 338 Proechimys steerei, 336 Profelis temmincki, 527 Prometheomys schaposchinikowi, 276 Pronghorn, 595 Pronolagus rupestris, 346 Propithecus tattersalli, 114 Propithecus verreauxi coquereli, 114 Prosomians, 103 Pseudocheiridae, 9, 38 Pseudocheirus peregrinus, 38, 59 Pseudochirulus cinereus, 38 Pseudochirulus herbertensis, 38 Pseudois nayaur szechuanensis, 656 Pteronotus gymnonotus, 398 Pteronotus macleayii, 398 Pteronotus parnelli, 397 Pteronotus personatus, 397 Pteropus giganteus, 384 Pteropus lylei, 384

711

Pteropus rodricensis, 383 Puma, 522 Puma concolor, 522 Pygathrix nemaeus, 157 Q Quagga, 672 Quoll: eastern, 29; northern, 28; spotted-tail, 30 R Rabbit: black-tailed jack, 353; bushman, 347; domestic, 354, 355; jack, 354; marsh, 350; old world, 354, 355; pygmy, 349; red, 346; swamp, 350; volcano, 348 Raccoon, 482, 483 Rangifer tarandus, 593 Raphicerus campestris, 636 Rasse, 502 Rat: African grass, 302, 303, 304, 305, 306; African marsh, 300; Amazon bamboo, 340; black, 281; blackclawed brush-furred, 297; brown, 282; brush-furred, 299; buff-bellied, 283; cape mole, 319; chinchilla, 334; common mole, 320, 321; dune mole, 319; Ehthiopian narrow-headed, 294, 296; elegant rice, 192; Florida water, 274; foothill rice, 190; French Guiana brush-tailed, 339; giant mole, 320; gray-tailed narrow-headed, 295; Guinear multimammate, 289; house, 281; Huallaga spiny, 335; kangaroo, 187; Kollmansperger’s multiammate, 292; Kusu, 302, 303, 304, 305, 306; long-tailed giant, 287; Luzor striped, 285; Mt. Isaroq shrew, 285; multimammate, 288, 290, 291; Norway, 282; pack, 188; pest, 286; Philippine, 284; rice, 192; rock, 333; roof, 281; shaggy swamp, 300, 301; short-tailed bandicoot, 286; short-tailed brush-furred, 298; Simon’s spiny, 337; slender-tailed cloud, 284; small-eared pygmy rice, 193; Sterre’s spiny, 336; Sulawesian shrew, 284; terrestrial spiny, 338; trade, 188; tuff-tailed spiny tree, 339; unstriped grass, 302, 303, 304, 305, 306; Viscacha, 330, 331; water, 194; West African shaggy, 301; wood, 188; yellow pygmy rice, 193 Ratel, 497 Rattus flavipectus, 283 Rattus norvegicus, 176, 282 Rattus rattus, 176, 281 Reindeer: Siberian, 593 Rhabdomys pumilio, 176 Rhebok, 621 Rhinoceros: black, 679; Indian, 677; southern white, 678; square-lipped, 678 Rhinoceros sondaicus, 661, 663 Rhinoceros unicornis, 663, 677 Rhinocerotidae, 663 Rhinolophus ferrumequinum, 393 Rhinolophus rouxi, 392 Rhinophylla fischerae, 416 Rhinophylla pumilio, 415 Rhinopoma hardwickei, 389 Rhinopoma microphyllum, 389 Rhyncholestes raphanurus, 19 Ringtail, 481

712

INDEX

Rodentia, 173 Romerolagus diazi, 348 Rousettus aegyptiacus, 381 Rousettus angolensis, 382 Rousettus lanosus, 382 Rousettus leschenaulti, 381 S Sable, 493 Saguinus imperator, 132 Saguinus midas, 133 Saimiri sciureus, 102, 131 Saki: white-faced, 123 Salpingotus crassicauda, 173 Sarcophilus harrisii, 30 Sassabie, 631 Scandentia, 95 Scotonycteris ophiodon, 386 Scotophilus heathi, 439 Scotophilus kuhlii, 438 Scotorepens balstoni, 436 Scotozous dormeri, 434 Sea lion: California, 531; northern, 531; Steller, 531 Seal: bearded, 534; common, 537, 538; gray, 535; harbor, 537, 538; harp, 535; Hawaiian monk, 532; hooded, 534; northern elephant, 533; northern fur, 530; ribbon, 536; ringed, 536; South African fur, 530; southern fur, 530; Weddell, 533 Seladang, 609 Serow, 649 Serval, 528 Sheep: blue, 656; domestic, 658, 659; European, 657; Rocky Mountain bighorn, 660 Shrew: Dsinezumi, 376; Elliot’s, 371; Eurasian, 368; Eurasian pygmy, 366; Horsfield’s, 373; Lagranha, 369; Laxmann’s, 367; lesser, 373; long-tailed, 366, 367, 368, 369; musk, 375; northern tree, 97; old world water, 372; pale gray, 374; Philippine wood, 360; pygmy, 375; Radde’s, 367; short-eared elephant, 70; Siberian, 374; southern short-tailed, 370, 371; whitetoothed, 373, 374, 376 Siamang, 160 Sicista betulina, 173 Sicista napaea, 187 Sifaka: Coquerel’s, 114; golden-crowned, 114 Sika: Dybowski’s, 587; Mandarin, 588 Sirenia, 71 Skunk: spotted, 499 Sloth: Hoffman’s, 84; two-toed tree, 83 Sminthopsis crassicaudata, 25 Solenodon: Hispaniolan, 366 Solenodon paradoxus, 357, 358, 366 Solenodontidae, 357 Sorex araneus, 368 Sorex caecutiens, 367 Sorex granarius, 369 Sorex minutus, 366 Sorex raddei, 367 Sorex tundrensis, 369

Soricidae, 358 Spalacopus cyanus, 332 Spalax ehrenbergi, 222 Speothos venaticus, 464 Spermophilus alaschanicus, 178, 180 Spermophilus erythrogenys, 178 Spermophilus undulatus, 179 Sphaeronycteris toxophyllum, 426 Spilogale gracilis latifrons, 499 Spilogale putorius interrupta, 499 Springbok, 648 Springbuck, 648 Springhaas, 310 Springhare, 310 Squirrel: chestnut great flying, 181; ground, 178, 179; Sunda tree, 177 Steenbok, 636 Stenella attenuata, 550 Stenella clymene, 545, 549 Stenella dubia, 551 Stenocephalemys albipes, 296 Stenocephalemys albocaudata, 294 Stenocephalemys griseicauda, 295 Stilodipus telum, 173 Stoat, 484 Sturnira erythromos, 417 Sturnira lilium, 416 Sturnira mordax, 417 Suidae, 564 Sulawesi babirusa, 571 Suncus murinus, 375 Sundasciurus philippinensis, 177 Sus barbatus, 568 Sus cebifrons, 568 Sus scrofa domestica, 566, 567 Sus scrofa scrofa, 567 Susliks, 178 Sylvilagus aquaticus, 350 Sylvilagus audubonii, 352 Sylvilagus floridanus, 351 Sylvilagus nuttallii, 352 Sylvilagus palustris, 350 Sylvilagus transitionalis, 351 Synaptomys cooperi, 275 Syncerus caffer, 605 T Tachyglossus aculeatus, 3, 6 Tachyoryctes macrocephalus, 223 Tachyoryctes splendens, 224 Takin, 651 Talapoin, 149 Talpa altaica, 377 Talpa europaea europaea, 376 Talpidae, 358 Tamandua, 86 tamandua longicaudata, 86 Tamandua tetradactyla, 86 Tamarin: black lion, 139; emperor, 132; golden lion, 139; red-handed, 133

INDEX Tamias sibiricus, 177 Taphozous longimanus, 390 Taphozous melanopogon, 390 Taphozous nudiventris, 390 Taphozous saccolaimus, 391 Tapir: Baird’s, 675; lowland, 673; Malayan, 676; mountain, 674 Tapiridae, 663 Tapirus bairdii, 664, 675 Tapirus indicus, 663, 676 Tapirus pinchaque, 663, 664, 674 Tapirus terrestris, 663, 664, 673 Tarsier: Philippine, 117; western, 116 Tarsipedidae, 9, 39 Tarsipes rostratus, 61 Tarsius bancanus, 116 Tarsius syrichta, 117 Tasmanian devil, 30 Tatera nigricauda, 222 Taterillus arenarius, 227 Taterillus congicus, 228 Taterillus petteri, 226 Taterillus pygargus, 226 Taterillus sp., 229 Taterillus tranieri, 227 Taurotragus derbianus, 600 Taurotragus oryx pattersonianus, 599 Tayassu pecari, 572 Tayassu tajacu, 573 Tayassuidae, 564 Tayra, 495 Tenrec: long-tailed, 68; streaked, 68 Thomomys bottae pervagus, 173 Thomomys talpoides, 184 Thomomys talpoides attenuatus, 184 Thomomys talpoides pygmaeus, 185 Thryonomys swinderianus, 174 Thylacinidae, 21 Thylacinus cynocephalus, 21 Thylamys elegans, 13 Thylacomys logotis, 32 Thylogale billardierii, 45 Thylogale thetis, 46 Tiger: Siberian, 519 Tokudaia osimensis osimensis, 175 Tolypeutes matacus, 92 Tonatia bidens, 379, 408 Tonatia schultzi, 379 Topi, 632 Toros, 339 Trachops cirrhosus, 405 Trachypithecus cristatus, 102, 158 Trachypithecus obscurus, 156 Tragelaphus angasii, 596 Tragelaphus eurycerus, 599 Tragelaphus imberbis, 598 Tragelaphus strepsiceros, 597 Tragulidae, 564 Tragulus javanicus, 576 Tremarctos ornatus, 447, 471, 472

713

Trichechus manatus latirostris, 72, 73 Trichosurus vulpecula, 37, 43 Trichys fasciculata, 174 Trichys lipura, 174 Tscherskia triton, 215 Tubulidentata, 80 Tuco-tuco, 325, 326, 327, 328 Tupaia belangeri, 95, 97 Tupaia glis, 95 Tupaia montana, 95 Tupaia palawanensis, 95 Tupaiidae, 95 Tur: east Caucasian, 654 Tursiops truncatus, 545, 552, 553 Tylonycteris pachypus, 435 Tylonycteris robustula, 436 Tympanoctomys barrerae, 174, 176, 330, 331 U Uakari: red, 124 Urocyon cinereoargenteus, 457 Uroderma magnirostrum, 417 Ursidae, 447 Ursus americanus, 474 Ursus arctos, 475 Ursus malayanus, 477 Ursus maritimus, 476 Ursus thibetanus, 473 Ursus ursinus, 478 V Vampyressa brocki, 419 Vampyressa nymphaea, 419 Vampyressa pusilla, 379, 419 Vampyrum spectrum, 405 Vespadelus sagitulla, 432 Vespadelus vulturnus, 432 Viscacha: plains, 322 Viverricula indica, 502 Viverridae, 449 Vole: Afghan, 245; Amargosa, 271; bank, 231, 232, 233, 234, 235, 241; beach, 270; Bucharian, 246; California, 271; Caucasian snow, 244; common, 258; Coronation Island, 254; creeping, 271; Daghestan pine, 249; European pine, 248; European snow, 242; European water, 241; Evoronsk, 267; field, 269; gray red-backed, 231; Günther’s, 256, 257; heather, 240; Japanese grass, 262; juniper, 247; large-eared, 239; long-tailed, 254; Major’s pine, 250; Maximowicz’s, 266; Mexicanus, 272; Mongolian, 264; Muisk, 266; narrow-headed, 268; Nasarov’s, 251; north Siberian, 264; northern, 277; northern red-backed, 233; Pere David’s, 237; prairie, 253; Pratt’s, 237; red-backed, 231, 232, 235; reed, 263; Robert’s snow, 243; royal, 237; Sakhalin, 265; Schelkovnikov’s pine, 252; Schiddlovski’s social, 256; Sikotan, 232; silver high mountain, 238; social, 255; southern, 259, 278; southern red-backed, 234; Sylvestre pine, 253; Tien Shan, 261; Tien Shan red-backed, 236; Townsend’s, 270; Transcaspian, 260; transcaucasian, 279; tundra,

714

INDEX

262; water, 254; western red-backed, 234; woodland, 253 Vombatidae, 9, 39 Vombatus ursinus, 41 Vormela peregusna, 492 Vulpes corsac, 453 Vulpes macrotis, 453 Vulpes vulpes, 451, 452 W Wallabia bicolor, 37, 50 Wallaby: black-footed rock, 47; brush-tailed rock, 48; little rock, 49; parma, 51; red-necked, 51; short-eared rock, 49; spectacled hare, 37; swamp, 37, 50; tammar, 37, 52, 53; whiptail, 54; yellow-footed rock, 46, 47 Wallaroo, 37; common, 57 Walrus, 532 Wapiti, 586, 591; Bactrian, 590 Warthog: southern, 571 Waterbuck: Defassa, 617; Ellipsen, 617 Weasel: Japanese, 488; least, 485; mountain, 486; Siberian, 487 Whale: Beluga, 548; blackfish, 555; blue, 562; bowhead, 559; fin, 560; Gervais’ beaked, 556; gray, 559;

Greenland right, 559; Hubbs’ beaked, 555; humpback, 563; killer, 554; minke, 560; pilot, 555; pygmy sperm, 557; Sei, 561; sperm, 558; white, 548 Wildebeest: black, 634 Wisent, 611 Wolf: grey, 466; maned, 469 Wolverine, 497 Wombat: common, 41; northern hairy-nosed, 42; southern hairy-nosed, 39 X Xenarthra, 81 Y Yak, 610 Yapok, 10 Z Zaedyus pichiy, 82, 90 Zaglossus bruijni, 4, 7 Zalophus californianus, 531 Zapus hudsonius, 173 Zebra: Burchell’s, 671; Grant’s, 672 Ziphiidae, 546 Zorilla, 496

The Orders and Species of Mammals Super-Orders Orders Monotremata Order Monotremata

*Total Number of Species

Number of Species with Karyotypes

Number of Ideograms

5

3

1

5

Marsupialia Order Didelphimorhia Order Paucituberculata Order Microbitheria Order Dasyuromorphia Order Peramelemorphia Order Notoryctemophia Order Diprotodontia

331

Afrotheria Order Afrosoridida Order Macroscelidea Order Sirenia Order Proboscidea Order Hyracoidea Order Tubulidentata

79

Xenarthra Order Xenarthra

31

55 87 6 1 71 21 2 143 15

2767

14

Laurasiatheria Order Eulipotyphla Order Chiroptera Order Carnivora Order Pholidota Order Cetartiodactyla Cetacea Artiodactyla Order Perissodactyla

2203

TOTALS

5416

303

425

21 110 15 815

0 0 1 0 0 0

63 63 63 71 74 78 80

0

81 81

0 0 12 6 1

95 95 98 100 173 342

1 2 11 0 6 1 5 1

357 357 378 445 539 541 541 564 661

21 27 140 110 1 131

84 240 17

9 10 19 20 21 31 35 36

19 1 1 77 209 15

452 1116 286 8 324

1 0 0 0 0 0 0

0 14

20 2 376 2277 92

1 1

1 9 1 1 2 1 1

31

1 1

10 1 1 10 3 1 29

51 15 5 3 4 1

Euarchontoglires Order Scandentia Order Dermoptera Order Primates Order Rodentia Order Lagomorpha

*Wilson & Reeder 2005

3

43

Page

Microbiotheria Peramelemorphia Notorycetemorphia Dasyuromorphia Diprotodontia

Australidelphia

Proboscoidea Hyracoidea Sirenia Tubulidentata Macroscelidae Afrosoricida

Afrotheria

Xenarthra

Xenarthra

Primates Scandentia Dermoptera Lagomorpha Rodentia

Euarchontoglires

Pholidota Carnivora Perissodactyla Cetartiodactyla Chiroptera Eulipotyphla

Laurasiatheria

Eurtheria

Ameridelphia Marsupialia

Major Clades Didelphimorphia Paucituberculata

Monotremata Cretaceous 200

125

100

90

Cenozoic 80

70

60

50

40

30

20

10

0

KT Boundary Million Years Before Present

An evolutionary tree depicts historic divergence relationships among the living orders of mammals. The Atlas presents available species’ karyotypes from each of these orders. The phylogenetic hierarchy is a consensus view of several decades of molecular genetic, morphological, and fossil inference. Details of molecular results and estimated dates can be found in the following references: (Amrine-Madsen et al., 2003; Murphy et al., 2001a, 2001b; Lu et al., 2001; Springer et al., 2003; 2004; Nilsson et al., 2003; 2004).