berlinerpaliiobiologischeabhandlungen, band 5
FOSSI DIBRANCHIATE CEPHALOPODS A PALE OZOOLOGICAL MONOGRAPH by
ADOLF NAEF 1922 with a frontispieceand 101 textfigures Translatedfrom German(original title: "Die fossilenTintenfische" l92z)
Edited by Kerstin Warnke,Helmut Keupp & Sigurdvon Boletzky Weinert GmbH Berlin 2004
DIE TOSSILEN
TINTEI\FISCHE EINE PALAOZOOLOCISCHE MONOORAPHIE VON
Dn. ADOLF NAEF PRTVATDOZENTEN FORzoolootE AN DERuNIvrRstTliT z0RIcH
MIT AINNM TITELBILD UND 101 ABBILDUNCEN IM TEXT
W JENA VERLAGVON GUSTAVFISCHER 1922
\ n'ord from the editors l:nglishtranslations of Adolf Naef s monographon the Cephalopoda of the Bay of Naples(1921-1928)havebeenmade .rrailableby the SmithsonianInstitutionLibraries,Washington,D. C., (1912,2000). The presentissueof Berliner I'alilobiologischeAbhandlungenprovides the English translation of the companion volume on fossil coleoid drbranchiate) cephalopods, which is ofinterest to both paleontologistsand neontologists. Careful proof-reading by Professor Desmond Donovan (London) significantly improved the preliminary :ranslationby S. v. Boletzky.Ail the resultingeditorialnotesare given in squarebrackets[...]. Moreover, Professor Donovan brought to our attention a list of corrigenda assembledby Riegraf, Janssen& (1998)*. This list (reproducedbelow) is here combinedwith further correctionsgiven by Professor Schrnitt-Riegraf I)onovan. Since the insertion of figures in the presenttranslation altered the original line numbering, referencesto -.rplanations of figuresare indicatedin squarebrackets[Fig. ...].Likewise, in the only casewherea corrigendumrefers ro an original footnote,this is indicatedin squarebracketsfNote rr]. (Naefs footnotesare successivelynumberedand .rssembled as "Notes" at the end of the translatedtext): x*x**:k*i<*rr*rr*rr*****16*****rF*rF**r<*r<*r<*)k***********r.************************************r<*********tr
Corrigenda: page numbers refer to the original text; line numbers in parentheses:
f,
Page
Line
l8
(11):
Owen 1836shouldreadOwen 1832
18
(15):
Owen 1836shouldreadOwen 1832.andLeach1818shouldreadLeach1817.
48
(23):
Belemnosidaewas proposedby Wiltshire, 1869.
s2
(1s):
5)
())\.
B e l e m n i t easn o m a l u sS o w . , 1 8 2 9 ( v . 6 , p . 1 8 3 , P l . 5 9 , F i g . 2 ) s h o u l dr e a d BelopteraanontalaSow.,1829(u. 6, p. 183,Pl. 591,Fig. 2). "Bullen" is Bullen Newton, i.e. Newton is the surname (also in list of
1, i
\--
t.
references).
55
(35):
63
(27) fFig.22l: DingdennearMtnster shouldreadDingdennearOberhausen. (5) fFig.231: of Westphaliafrom Dingden and Bersenbrtickshould read of Rhinelandfrorn
64
Deshayes1824shouldreadDeshayes1837.
6s
(20):
68
(20):
Dingden near Oberhausenand Bersenbriick near Osnabnick, administrative areaof Hanover. of Westphalianear Dingden and Berssenbriickshould read of Rhineland from Dingden near Oberhausenand Bersenbriick near Osnabnick, administrative areaofHanover. Heliceras(Dana 1848)shouldreadHelicerus.. .
68
(21):
H eI i cer trs .firgerzszsshould r eadfu eg ensis .
70
(14):
The speciesnamespirula datesfrom Linnaeus1758.
76
( 1 5 - 1 6 )[ F i g . 2 e ] :di Fanganoshouldreaddi Fangario.
82
(2):
BelosepiaVoltz 1930shouldreadBelosaepiaVoitz, 1830.
82
(6):
Deshayes1825shouldreadDeshayes1837.
82
(10):
Belosepiacuvieri shouldreadBelosaepiacuvieri.
82
(11):
longirotris should readIongit"ostris.
e3
(12):
Rcimerand DamasshouldreadRoemer& Dames.
rOe
(33)
I 13
(23):
l 15
(1):
fNote
33]
SteiningenshouldreadHeiningen. Plesioteuthis was proposedby Wagner1859,not 1860. Acanthotetrthis tricarniata shouldread ... tricarinata.
L
I 15
( 19):
On.,-c'hoteuthis tricorniatushouldread.. .tricorinata.
I l9
(25):
LeptoteuthidaeshouldreadLeptoteuthididae.
I 19
(28):
Leptoreuthis H. v. Meyer shouldreadLeptothefihi.s...
122
(22-23):
Loligosepiai.vasproposedby Quenstedt1839,not 1843.
122
(26-27):
125
(26'):
125
(26):
BektpeltissintplexVoltz, 1840is not the type speciesof Geotettthisv. Mtinster, 1843: the type speciesis Loligo bollensisSchiiblerin v. Zieten, I 832. Loligo actlensisv. Zieten shouldreadBelopeltissinLtatusYoltz Loligo aalensis is an invalidly designatedtype species,becauseit is not quoted in the original description; Belopeltis sinuatus Voltz was subsequentlydesignatedtype speciesof Belopeltisby Doyle, Donovan& Nixon, 1994:9. Referencesto Zieten (Loligo aalensis)hereand elsewhereshouldbe 1832.
121
(29):
Trittlingen,Bo11,Mittelgau. Ahlen should read Frittlingen,Bo11,Mistelgau, Aalen.
132
(5):
TrittlingenshouldreadFrittlingen.
132
(21):
nurnber8764shouldread...8746.
134
(9):
134
Sahil Almae shouldreadsahilalmue.
(33-34) [Fig.49]: Sahel-Almaeshouldreadsahilolntae,
134
(40):
139
(2):
142
(26'):
143
(l).
Beloteuthi.s leckenshyishouldread .. .leckenbyi.
144
(3):
u p t o 5 0 c m ( Q u e n s t .1 8 5 8 ,p . 5 0 6 ) s h o u l dr e a du p t o 3 5 c m ( Q u e n s t .1, 8 5 6 -
Sdhil-Almaeshouldreadsahilalntae. in the explanation oftextfig. 52, I 866 shouldread1896. Lolig ohrevisBlainr'.shouldreadLoligo bret'is...
I 8 5 7 ,p . 2 4 3 . 148 149
( 1 a ) [ F i g .5 5 ] : o h l r t r t g(uW a g n e r1 8 4 8 )s h o u l dr e a d. . . 1 8 5 9 ) . (26):
Mahn EpsilonshouldreadMahn Zeta.
172
(44) fFig. 651: Conoteuthis dupianusshouldrcad...dupinicutu.s.
112
(aa) [Fig 65]: explanations k and I appearto havebeentransposed. of iter.r.rs
117
(26):
1 8 3 2s h o u l dr e a d1 8 3 9 .
119
(30):
Lyma-RegisshouldreadLyme Regis,Dorset,England.
185 196
(23) [Fig. 66]: Belemnites brughierishoLrld read.. .bruguierranus. (10):
B. brughieriMiller shouldread...bruguierianus d'Orbigny.
205
(25) fFig.711: P. (A.) ohltnannensis shouldrcad...ahlumensrs.
205
(?) enigmoticzrs (aa) [Fig.71]: Doc4'loretrrhis shouldread...aenigmatictts.
205
(47) [Fig. 7l]:
Bt'ac'h.t'teuthis shouldreadBrachl,belus.
225
( 5 6 ) [ F i g . 7 l ] : d ' O r b i g n y 1 8 8 6s h o u l dr e a d . . . 1 8 4 5 ;t h e d r a w i n gi n F i g . 7 1 v i s a c o a r s e l y represented simplebelernnitealveolus. (27'): BelemnitesclavatusStahl,1824is the type speciesof Hastite,s.not: Belemnite,s clat'atusv. Schlotheim,1820. (15): Raphibeltrs shouldreadRhaphibelus.
225
(17):
225
(18):
225
(18):
225
(19):
205 224
BelemnitessulcatusMiller, 1826 is the type speciesof Belemnop.slsBayle, l878, not Belemnites bessinus d'Orbigny,1842,as it is maintained. Hibolites Mayer shouldreadHibolithesDenysde Montfort. Hibolithes hastatusDenys de Montfort, 1808 is the type speciesof Hibolithes (by monotypy),not Belentnite.s ha.statusde Blainville, 1825, as erroneously stated. B e l e m n o c o n u(s. b a u d o u i ndl ' O r b . ) s h o u l d r e a d B e l e m n o c o n u s n o v . g e n . ( b a u d o u i ndi ' O r b . ) .
3 225
(28):
226
(19):
PseudoduvaliaQtol,vgonalisBlainv.) should read Pseudoduvalia nov. gen. (polygonal is Blainv.). Rhobalobeltrs shouldreadRhopalobelus.
234
(22):
88c shouldread 88e.
245
(1):
249
(12):
249
(27):
249
(28):
O. jasikovishouldreadO.jasikowi. BelemnitessulcatusMiller, 1826 is the type speciesof BelemnopslsBayle, I 878,not Belemnitesbessinusd'Orbigny, 1842,as eroneously stated. Hibolites Mayer-Eymar1883should readHibolithesDenysde Montfort, 1808.
257
Hibolithes hastatutsDenys de Montfort, 1808 is the type speciesof Hibolithes, not Belemniteshastatusde Blainville, 1825,as erroneouslyquoted. (21) [Fig.93]: Hastitessouyananishouldread...sauyanausus.
259
(2):
B. gilieroni shouid read...gillieroni.
259
(9):
C. ertinctoriusshouldread C. exstinctorius.
259
(29) lFig.9al:
BelentnitesrltgosusshouldreadBelemnitesrugfbr.
280
(12):
LauffenbergshouldreadLaufenburg,Baden(on the river Rhine).
280
(18):
Pageref. to Systemshouldbe 534.
294
(27):
johanneusHilter shouldread ...Hilber.
294
(34):
BoenninghausshouldreadHoeninghaus.
299
(12):
SepiodeashouldreadSepioidea.
**********tl:------tll]-**********l*t:'_'::_Ti.'_."_"1:::::::___'_'::::::_______****{.,!,rx*,F****,< Only the most obviousmisprints(e.g.Lolig obrevis)were "silently" comectedin the translatedtext. Page referencesin the text refer to Naefs original pagination,which is throughoutgiven in parentheses, in bold italics. An important changefrom the original book is the presentsize of the type area.A11Ihe/igures are enlarged 1.4 tiraes, thereforethe size indicationsgiven by Naef in his figure explanationsmust be changed(".g. "t/. natural size" becomes0.7 nat. size).We deliberatelyrefrainedfrom making this changein the new text. Although Naef s book appearedmore than 80 years ago, we had to ask permissionto publish this translation.We thank Urban & Fischer(the successorof GustavFischerVerlag, I 1.04.01)for grantingthis permission. An entirely new appendixis added,containinga list of synonyms,whtch was kindly provided by Dr. Theo Engeser (FU Berlin). We thank Petra GroBkopf (FU Berlin) for technical assistancewith the illustrations and Micha Bustian for secretarialhelp. We are most gratefulto ProfessorDesmondDonovan for all his encouragement and invaluablesupportthroughout the preparationof this translation. Kerstin Warnke,Helmut Keupp & Sigurdv. Boletzky
*) Fossilium Catalogus I: Animalia (Ed. F. Westphal) Pars 135: W. Riegraf, N. Janssen& C. SchmitrRiegral-.A. Cephalopoda dibranchiatafossiles(Coleoidea)II [incl. Addenda et corrigendaad Fossilium Catalogi Pars I 1. Cephalopodadibranchiata(1920); Pars 130, Vampyromorpha(1988); Pars 133, Cephalopodadibranchiatafossiles(1995) et ad A. Naef. Dic fossilenTintenfische.Jcna (1922)1,pp.25-27.BackhuysPublishers, Leiden,1998.
Table of contents German English page
page
A rvord from the editors - K. Warnke, H. Keupp, S. v. Boletzky
I
Preface- A. Naef
f
Introduction
(1)
6
Part I: Technicalprerequisites
(11)
e
Part II: The Sepioidea,or Sepiu-likecoleoids
(38)
24
Part III: The Teuthoidea,or squid-like coleoids
Q01)
62
Part IV: The Belemnoideao or belemnite-likecoleoids
(164)
ee
Part V: The Octopoda,or octopusJikecoleoids
(283)
16s
Concluding remarks and systematicsynopsis
(2es)
r72
Notes
179
Referencelist
(s04)
Index
(sr7) re7
List of new species
(322)
List of synonyms- T. Engeser
(Contentsare given in greaterdetail at the beginningofeach major section)
187 202 203
FOSSIL DIBRANCHIATE CEPHALOPODS A PALAEOZOOLOGICAL
MONOGRAPH
by ADOLFNAEF 1922
Preface. This review of fossil dibranchiatecephalopodsis completein itself. lt attemptsto presenta greatvariety of old and nelr' tnctsin their natural context,as far as is now possible.At the sarnetime, it is also part of a greaterwork, a synopsisof c.-phalopodsin general,comprisingboth living and extinct representatives of the class.Within this generalframework, this book is the counterparlof a monographof the living cephalopodsof the MediteraneanSear. Being part of this greaterwork, the presentbook also has a general aim, namely a comprehensiveexpositionof the problent of organic diversity, and especiallyan analysisof the prerequisites,the methodologyand the range of applicationsof the historical approachto this problem. To reachthis goal was conceivableto me only through intensive researchwithin a carefully chosen framework, starting from the living representativesof the group: they had to be analysedin terms of their systematics,comparativeanatomy,embryology,physiology,and life style in relationto living conditions, before their extinct relatives could be given a synoptic treatment.Together these two lines of research should lead to very generalconclusions,thus assistingin the scientific consolidationof evolutionarytheory basedon new, comprehensiveand exhaustiveinsights gained from personalwork. The need for this approachhas alreadybeen p o i n t e d o u t i n m y e a r l i e r p u b l i c a t i o n s ( c f . N a1e9f1 1 ,1 9 1 3 ,1 9 1 7 , 1 9 1 9 , 1 9 2 0 , 1 9 2 1 ) , p e r h a p s w i t h o u t m a k i n g m u c h impressionon scientific circles that are either not interestedin, or persist in traditional views on, morphology. Zoologistsof both schoolswill find more factsthan "speculations"in this book; but the very natureof our work requires a clearexpositionof the ultimategoal. I am indebtedto rnany people for their support of my palaeozoologicalstudies.First of all I have to thank the Scientific ResearchFoundationof the University of Zurich and their generouspatrons for travel grants. I gratefully acknowledgethe directorsof foreign museumcollections,especiallyProf. Dr. F. Broili in Munich (BavarianState Collections)for his very liberal supporl.The hospitality of the museumsof Stuttgart,Tiibingen,Vienna and Berlin, and of the special collectionsat Holzmaden (Mr B. Hauff , Eichstiitt (Prof. Schwertschlager)and Braunschweig(Prof. E. Stolley)is also gratefullyackrowledged.
0) Introduction.
W e i n d e e d p o s s e s sa n e n o r m o u s a m o u n t o f literature, the research aim of which is termed "phylogeny" (i.e. history of phyla), but which deals
Although we are going to deal here with facts,and thus
exclusively with extant forms. We have textbooksand
sparethe opponentsof methodologicalexcursionsany
handbooksof systematiczoology and botany explicitly
(ostensibly)steriledigressions,neverthelesswe have to
based on "phylogenetic" considerations,in which
deal with some general consideratiorzsfirst, because
fossils are rarely or never mentioned. Renowned
without them an understandingof the facts would be
zoologistshave attemptedto reconstructthe historical
difficult. Without continuins reflexion. no sciencewill
"development"of the animal kingdom and discussed,
everthrive!
with great wisdom, whether the molluscs are derived
We are going to deal with an animal group that is a
from annelids or platodes, whether the trochophora
diversity of typically similar organisms representedby
larva of extantannelidsis a primitive form, whetherthe
(generally extinct) fossil forms, and attempt to follow
lining of the blood vessels is derived from a
them through geological time in their natural
coelothelium(somethingnever observedin embryonic
systematiccontext.Individual fossils will be viewed
development),etc. ln general thrs strange historicism
from an explicitly biological perspective,in other
has taken very little notice of the fossil record of the
words as the product or part of a living whole, which
past.
will be treated as a living animal, as far as
On the other hand,palaeontologlsls have lined up
permit. We considerthis the essenceof a circumstances
shellsupon shells,and seriesof bones,working in their
"palaeobiological"(sensu lato) approach,which must
turn on "phylogeny",without paying much attentionto
remainin contactwith the actualisticlife sciences.
the knowledge gained by their zoological colleagues.
Unfortunately,all we have at our disposalto begin
Thus bridges are constructedthat span millions of
with are corpses, in general grossly maltreatedand
years,with the convictionthat they have solid scientific
incompletecorpses.Most of the time there are only
justiflcation.
fragments,generallyparts of a skeleton.Palaeozoology
In contrastto these tendencies,one should realize
can rarely base its analyseson other traces of lile in
that in generalneitherthe morphologyof living beings,
past environrrents(tracks, burrows, etc.). Thus the
nor the rnorphologyand stratigraphyof fossil forms are
b a s i c c h a r a c t e r i s t i co f t h i s d i s c i p l i n e w i t h i n t h e
able to clarify the real pattern of descent among
framework of biology is clearly defined by the nature
organic species2. (J/
of the objects studied. Palaeontologyis an essentially
combination of these two disciplines, which have
morphological disciplineand can only be significant
become separatedby an ar1ificialdivision of labour, is
within the framework of such a scientific endeavour
necessary.Whether this combinationwill really allow
(cf. Naef, 1917, p. 69) (2) Experimental treatment of
us to succeedin reconstructingthe history of phyla,
the subject is indeed ruled out, and a strictly causal
and under what conditions this is possible, are
analysisis thereforeinconceivable.
questionsthat remainto be examined.
But it is evident that a
logical
The first questionin fact is whether this really is a
considerations:the generalquestionsraisedby a fossil
scientific aim. Some modern biologists will readily
This
determines the
nature of
dwell in the domain of systematicmorphology and its
agree that phylogeny is indeed in the state depicted
"comparative"methodology(Where does it belong?
here in a deliberately extreme manner. But they may
Where does it come from?). Within this domain,
considerthese problems to be concernsof the past,
however,palaeozoologyplays a leadingrole. Indeed,
which do not deservefurtherdiscussion.
palaeozoologyis indispensable for it (cf. Naef, 1919,
It is true that modernzoology has turned away from
pp. 28 and 56), especiallyif this disciplineis pursued
phylogeny,by which it has been disappointed,leaving
i n c o n s i d e r a t i o n o f h i s t o r i c a l p e r s p e c t i v e s ,i . e .
the rnatter in the state where it happenedto be. The
according to the predominant,basic theoreticalviews
mainstream of research now moves along other
of our time. In this respect,however, a very peculiar
pathwaysand forgets the stimulation and foundationit
situationpersistsin present-dayscience:
received frorn approacheswhich have since been
.rbandoned.
inevitable.But I refuse to believe that it generatedthe
In contrastto this trend, it should be recalled that \\'stematic morphology - which has always been the
extraordinary broadening of our detailed knowledge
cssenceofphylogeny
is the backboneand the root of
responsible for the increasing superficiality of
cr en the most modern trends in biology, and the
phylogeneticstudies.Here it must be made very clear
during the past 60 years; on the contrary, I considerit
liamework of the scientific study of life. ln this sense
that I am absolutely not opposed Io Ihe theorlt of
rr c contendthat: eachorganismhas two aspects,one is \ t t'ucture (form), the other is ef/'ect (function).
ultimate destination.But I considerthe theory to be
However, function is essentially,perhaps entirely
incompleteand in needof further development.A more
.lererminedby the given "form", in other words by the
profound developmentof systematicmorphology may
rpparatust Although the latter is also a product of an
further improve knowledge of phylogeny, which
cttect, that effect in its turn was realizedby an earlier
actually depends on morphology.
structure.Therelore morphology has implicit printac'y
commit the error of believing that searching for
in the frameworkof biology: Form can only result from
"relationshipsof form" and for ancestorsis one and the
tbrm, but it is forrr that also permits anything else.
samething!
heredity. That theory was our startingpoint and is our
One should not
This is also true of the inorganic realm. (The speci/ic
Since phylogenetic relations always remaln
lcatureof any effect is the consequence of a structure!).
hypothetical, and since one cannot deal with a whole
Systematicmorphology deals in particular with the
scientific domain exclusively in subjunctive form,
problem of natural order, consideringthe diversitv of
phylogeneticwork has adopteda deliberately explicit
organic forms and their correspondingachievements
way of thinking, which is not only questionablein (5,)
rrnd relationships.This problem is also approached
ideological terms but often has not donejustice to the
through phylogeny,but its solution has been sought in
particularneedsand the subtletyof scientificproblems.
l precipitate,arbitrary manner.A seriousconsideration
In contrast, the abstract way of thinking which is
of the whole state of affairs leads to the following conclusion:(4) there is no direct accessto blood
typical of idealistic morphology imposesvery careful,
rclationships and real genealogy from systematic
circumspectreasoningin the use of isolated facts and deters incompetentpeople from entering the field - a
s t u d i e s , e i t h e r b y n e o - m o r p h o l o g i c a lo r p a l a e o -
welcome consequenceindeed.It may then happenthat
niorphological methods. Rare fortunate combinations
the indirect approachyields more information on actual
confirm the rule. Thesedisciplinescan not thereforebe
developmentsthan could be obtainedin the customary
directedaccordingly,in other words they cannot be
way.
primarily focused on the special questionsof descent Attempts to do so have led [Ger. Abstammungsfragen]. to the logically absurdsituationalludedto earlier. What we can securely know about the origin of
In our field of research,the usual logical and conceptualtools are above all the units of the natural system(species,genus,family, etc.)r.But theseare not able by themselves(through their diagnoses)to fulfill
organic speciesis always very inadequatewith respect
the task. What is needed here is a special kind of
to the task of systernaticmorphology. The only
reasoning(cf. Naef, 1919)that has to be developedin a
solutionto the problem is to return to the firm basis of
"theory of typical similarities".A comparisonof the
classical (idealistic) morphology and embark on a
diverse forms of a group must be establishedon the
search for the "t1,pical similarities" [Ger. typische Ahnlichkeitenl of living beings (in terms of
basis of norms [standards,rules] or "types", i.e. the imagined wholes, ideal forms, that have proved
relationships of form) [Ger. Formverwandtschaften]. (Seethe Introductionto the "Cephalopoda").This may
extremelyuseful in solving the practicaland theoretical tasks of natural systematics.They are indispensableas
look like a reactionaryattitude, and in a senseit is a
methodologicaltools, and their application lies at the
reaction; it is indeed a fundamental rejection of the
core of a specialmethodologyin systematicbiology.
trend in biology that was inauguratedand invented by
Whether or not the types are subsequently
Haeckel, which appearsas an emor in methodological
interpreted as ancestralforms, they occupy a clearly
terms. It may be historically and psychologically
definedposition in the picture of organic diversity.But
understandable,and it thus may have been virtually
there is an important difference between the two
approachesmentionedearlier:namely,whethersuch an
Types thus form the most valuabletool for special
interpretationis hypothetically anticipatedor testeda
morphology, especially for palaeomorphology. As
posteriori. At any rate,we must refutethe view that the
mentioned earlier, the latter indeed suffers from the
recognition and representationof typical relationships,
fragmentary nature of the available material. For
i.e. the whole approachof our idealisticmorphology
biological employmentof this material (p. 1), some
(which is not our invention, but has been more
reconstruction is almost always necessary. Such
precisely and acutely defined by us), is sirnply a
reconstruction can only be done on the basis of
restatementof Haeckelianphylogeneticsthat would
classicalmorphology, i.e. following Cuvier'sprinciple
better be retranslatedinto the latter (in the sense "typical - primitive; type : ancestor, etc."). The
o/ correlation. This principie of coursemust be refined within the framework of the methodology outlined
contraryis true, as has been shown earlier (Naef, 1919,
herein, leading to the following formulation: For the
p. 35). In addition to the historical priority and the
reconstrtrctionof fragmentary remains oJ'an animal,
methodologicaladvantagesdiscussedabove, the ideal
the missing elements o./ organisation have to be
(6) type also has absolutepriority over the hypothetical
assumedto correspond to the probable typical .features
ancestorfor generalbiologicalreasons.This becomes
of the respective group. This of course requires a
clear as soon as we considerlhe conceptof the ttornt itt
detailedknowledgeof the systematicmorphologyof all
biology, from which the type concept is ultimately
recent and fossil relatives; in particular knowledge of
deducible.
their naturai systemand its foundations,in other words
Characters are typical not only of systematic
of comparativeanatomy and developmentalhistory!
categories,but also of single living individuals.The
A scientific treatment of extinct organisms can
latter do not in fact have an inflexible configuration;
thereforeonly be achievedby specialistsfamiliar with
they show diverse, variable manifestations,which we
their extantrelatives5.
weigh against one another, consciously or
(8) The way in which special knowledge of
unconsciously,to reach a generalpicture or concept
relationshipsof form shouldbe usedin reconstructions
through abstractiona.ln systematicsthe concept oJ the
may be clarified by some additional indications: very
type is stretchedbeyond the ontogenyof an individual
often (Fig. 101) we have no living representativefor a
or the sequenceof generationsand is applied to the
restrictedgroup of fossil species;in such caseswe are
species as a natural unit (a reproductive comntunity)
obliged to refrain from consideringonly the special
and furthermore extended,step by step, to all beings
features of this group and have to include the next
connectedby "typical similarity" or relationshipof
broaderone. Perhapsthis will still be impossible,so we
form. The type concept thus becomesthe essential
have to proceed,step by step,to more remoteforms for
meansof expressingsuch relationshipsand provides
comparison.It may then happenthat theseforms do not
the conceptualframework for the mental assimilation
(any longer) have the relevanthomologousfeatures,so
of such an immensediversity. The particularformative
that we can only get some information from the
norms become the objects of comparisonand the raw
embrl,os by consideringtheir group-typical structure.
material for the study of the increasingly generalized
In this way we may find a basis for the reconstruction
norms, similar in a way to the "types" and "bauplans"
of e.g. the belemnites(see the systematicoverview):
of classicalmorphologists.
there are no living representativesfor the whole
What is typical can be recognized from a
suborderBelemnoidea.The closestliving relativesare
rnethodicalcomparison,the principles of which (Naef,
the Teuthoidea,in which we can at leaststudy the roles
1919,pp. 25-33) are derivedfrom the very nature of
of the pro-ostracum,the phragmocone,and the rostrum
the real objects,especially(7) from their real similarity
in terms of anatomy, developmentalphysiology and
c'onnexions[Ger. Ahnlichkeitsbeziehungen] that follow t h e i r o w n 1 a r . r . , sT. h e v e r y e x i s t e n c e o f t h e s e
ecology.Startingfrom thesedata,which can be viewed
relationshipsallows us to considerthe norms that we
the Sepioidea,we first reconstruct,backwardsas it
have recognizedas naturally inevitable definitions,
were, the decapodanorganisation,and then again
trom ri'hich definite and (as will be shown) successful
forwards, the organisation of the belemnoids and
cr)nclLlsions canbe drawn.
belemnites.
in parallel with those obtainedon the phragmoconeof
If palaeomorphology is a part of systematic morphology,by the other parts of which it needsto be
morphology and its correlates(systematicphysiology, systematic ecology, systematic biogeography,
supported, it provides in return a substantial
systematicbiostratigraphy). The above sentencedoes
cnrichmentof the whole. This enrichmentis due to two
not ofcourse cover all ofthe contentsand extentofthe
sources:extinct speciesvery substantially enrich the
modern theory of evolution, nor can it suffice as a
.stockoJ Jbrms (in the present group, the number of
methodologicalprerequisitefor (10) the future practice
extinct species is about 20 times that of living
of the discipline from which it has arisenT.But it is the
speciesl).Moreover, the fossils are dateable,as groups
dominant principle of modem biology underlying any
. in terms of geological levels; thus they translatethe
comprehensiveapproach; it is both interpreting and
natural order as expressed in the system into a
being interpreted.
historically ordered diversification of formative norms (which are usually viewed as [Ger. Gestaltungsnormen] groupings). species-genealogical One person may consider the system as directly .s.t'mbolizinghistory (a family tree), whereas another
(11)
Part I: Specialprerequisites
personwill view it as the natural,complex step-ladder, or.rwhich (9,) successivegenerationshave climbed upwards. The term "parallel evolution" refers to the progressof phyla as a common forward movement of great numbers of species each passing through a regular sequenceof [evolutionary]stages.This view is supportedby many observationsand is now almost predominant.In particular it is the generaltendencyof G. Steinmann and his school. Although we are
C o n t e n t s : A . O n s o m e b a s i c c o n c e p t so f m o l l u s c a n moryhology(p. 11).B. On the specialfeatures of ccphalopod (p. (p. shells 14).C. Diagnosis of the Class 17) D. On a systematic surveyof the cephalopods (p. 18). E. On the (p. distinctionbetwecntetrabranchiates and dibranchiates (p. l9). F. On the contrastbetweenoctopodsand dccapods 25). G. The suborderof decapods(p. 30). H. On the morphologyof the relationships betweenthe shell and the lins(p.35).
definitely opposed to it, we cannot a priori deny it somevalidity6. In our view, Steinmann'sideas provide an additional
reason for
A. Somebasicconceptsof molluscanmorphology
decouplingsystelnatic-
morphological problems from their phylogenetic
Before approachingour actual subject, we have to
interpretation.We may well imagine the ancestral
clarify some general ideas that will have to serve as
forms of our groups as somethingvery similar to the
prerequisites for any further understanding.
typesdepicted(Figs 11, 39, 62). I believe,indeed,that
The cephalopodsare ntolluscs and clearly exhibit
such a view will becomeestablished. But it is still a
the featuresof this phyletic relationshipin their entire
personal view and cannot yet be considered as
organisation,especially in the formation of the shell
scientifically corroborated knowledge. With this
and its relationship to the soft body. The following
qualification, neither the concentration
of
schematicrepresentation(Fig. 1) attemptsto illustrate
contparison on types, nor the
the general molluscan character in its typical form,
ntorphological
establishmentof their normative character(p. 6), nor
without claiming to be a secureresult of morphological
the basic view of evolutionarytheory are called in
abstractionin all the parts.Especiallythe problem of
question.The latter at presentstatesmerely this: in the
the primary subdivision of the headfoot (cf. Naef,
coltrse o/ geological historl, minor modiJicationsof
1 9 1 1 ,p . 8 3 ; 1 9 1 3 ,p . 3 8 5 )a n dt h e r e l a t i o n s h iw pith the
formative nornts have accumulated,by **hichprocess the present-davspecieshaye beengenerated.fromone
a m p h i n e u r a n s( N a e f , 1 9 1 1 , p . 7 7 ) s h o u l d n o t b e
or severalassumedancestralspecies.
simply note the generalfeaturesof the shell, which are
This hypothesis,which has long since become a
anticipated.We ratherlet the figure speakfor itself and to be consideredmorphologically printary compared
well establishedtheory, is independentof any opinion
with the more complex features often observed in
or doctrine about the specific process of species
cephalopods.
change; it is the general result of systematic
10 Fig. 1.
a) A diagramof molluscanorganisation.The f,igureis
a generaidiagram ofthe systematicallydefincd typical features of molluscs.There is some uncertaintyabout the overall aspect becauscthe Amphineura (Solenogastres,chitons) are diff-erent from the Eumalakia (all other classes).Thus it is uncertain whether the primary foot was as uniform as in the figure. whether it was fused w-rththe head.and whcther the shell was a tall coneor ratherflat. 1. cerebral;2. pleural;3. pedalpart ofthe circum-oesophageal rtng 4. statocyst;5. foot; 6. radiating musclesof the head-foot retractor: 7. base of head-fbot retractor; 8. anterior brachial (parietal) ganglion; 9. anterior limit of the mantle cavity roof; lri
13
10. posterior mantle cavity; 11. lateral free mantle margin, 12. same,postcrior, in optical median section,to show its relation to the ostracum (black). which grows by accretion here; -13. mantle fbld; 14. posterior: 15 anterior gilll. 16. typical position
30
of the anus at the posterior end of the mantle cavity roof; -17.
3r
visceralganglion;1B. heart;19. intestine;?0. shell epithelium; ?1. hypostracum(nacrcouslayers); 22. ostracum(porcellanous layer): x. apex (embryonic shell): 2J. stomachl2y'. shell muscle (head-fbot retractor); 25. fore-gut; 26. pleuro-visceral strand: 2/. most anterior point of mantle groove; 28. anterior mantle
cavity;29. eyc; -10.tonguein opticalsection;31.buccalcavity in opticalsection;32. snout;JJ. radularpouch;a. anterioraorta;/. liver; g. gonad;p. pericardium. 6) Sectionthroughthe mantle rnargin (13) with the fso-called]shell fold (r). The latter lies on the outsideofthe shell and frequently secretes,in addition to the typical shell layers(ostracumand hypostracum)which are formed by the primary shell epithelium (20), a third layer. the periostracum(z). When the shell fold is poorly developed.the periostracumis similar to the ostracum"in other words it is addedto the margin and increascsin thicknessdistally. It, howevcr, the shell fold covers the entire shel1,then the "secondary shell epithelium" can concentrateits function at any point: but it will always secretethe greatestnumber of layers at the apex. The periostracumis then more similar to the hypostracum.
The sheli is secretedby the "shell epithelium",i.e.
can generallybe tracedby foliowing the growth lines.
the part of the epidermislining the'lnantle sac", and it is therefore(12) wholly inaccurateto say that the shell
The gron-th in thickness is achieved by addition of
is "producedby the mantle". Its growth is the result of
of the shell, thus producing a "mother ol pearl
continuousaccretion,only exceptionallyare certain
(nacreous)layer" or "hypostracum";its thickness,in
parts subsequentlydissolved. The basic conchiolin
contrast to the afbre-mentioned layer, increases
IGer. "Conchin"] is an organic! tenacious, elastic substancethat hardensby dehydration,calcium being
towards the oldest parts of the shell, since they
subsequentlydepositedinto its fine layers,making the
thickening layers. Tl.reseof coursethin out completely
shellboth rigid and fragile.
towards the lnar-ginof the ostracurn.The hypostracum
The accretionof new shell materialis achievedin
foliate, thin layersof shell materialon the inner surface
n e c e s s a r i l yh a v e r e c e i v e d t h e g r e a t e s tn u m b e r o f
is producedbv the greaterpart of the shell epithelium,
I\\ro ways: namely as marginal growth and as growth in
whereas onl,v tl.relnarginal parts of the latter, which
rhic'kness.The former resultsin the enlargementof the outline and - accordingto the direction of accretion-
belong to the (prir.r.rary) mantle, are involved in the
deterrninesthe outer shapeof the shell, which may
A special strllcture and form is often found in the
clrangeconsiderably(13) in the courseof growth. It
oldcst parts of the shell, which are situatedat the apex.
producesa very solid shell layer the thicknessofwhich rncreasesfrom the t'apex"; this layer is called the "ostracurn"or "porcellanouslayer"; its development
must alreadvbc presentas a first rudiment.The earliest
formationoi the ostracum.
Indccd. belbre a sl.rellcan grow the way described,it cmbrvonic part of the shell therefore demands a
1l
Fig, 2.
A schematicrepresentation of cephalopodshe11 morphology.
.I) Prototype,Endoceras-ltke. The general molluscan she1lcomponentsare recognisable(cf. Fig. l): Apex (15), ostracum (7), (6.4, B,9. 10. 11,z, 13). The annulus(5. l2) andthe annulusmaterial(12,13) mark the divisionofthe hypostracum hvpostracum urto an outer (e) and an inner parl (l). Chamberformation (which doesnot start from the posteriorend!) is achievedby (first dorsally) \cparatingout thc componentlayersofthe inner part. The chamberlayersare diffcrentiated(with the exceptionofthe initial cap) into .t septum(B) and siphuncularcone [septalneck, the very first one in apicalposition] (v). In the latter,one can distinguisha calcareous conc (9) and a conchiolincone (11). Secretionsinsidethe chambersprovide the supporlingridges (.t) lbr thc septaand the pillars (10) tbr the septalnecks.1,fmarksthe] insertionofthc head-footretractoron the annulus. i,) Derived,normal form, Orthoceras-l1ke. The beginningof the siphunclehas separatedfrom the initial cap (:), with which it is only connectedby a strongpillar ("prosiphon", I 7). The gas chambers are now ventrally continuous, thus surrounding the siphuncle. which extends through the length of the phragmocone.The initial caecumofthe first siphuncularsection(18) is in generaluncalcified(in this respectNautilusis similar to rr). forming a conchiolincup; in the subsequent cups(19) suchinitial caecaareno longer distinguishable.
pafticular morphoiogical importance;it may in fact be
retractors;where they insefi on the shell wall, the shell
rather distinct from the subsequentparts. However, we
epithelium may be modified, so that the corresponding
do not considerthis "shell nucleus" to representthe
parts of the hypostracum also take on a special
(often wrongly) so-called"embryonic shells", which alreadyshow at leastpartly - typical charactersE.
character. This is particularly important for the
lf a separate"shell fold" fof the mantle] is formed extending over the outer shell surface - as often happens,producing a secondarycomplication-, then a
observedin other forms as well.
third shell layer can be added to those already
B. Specialfeatures ofcephalopod shells
interpretation of cephalopod shells, but it can be
mentioned;it may be called the "periostracum" or "outer layer" (Fig. 1b). Much like the ostracum(from
A characteristicfeature of cephalopodshells rs an
which it has become separated)it will increasern
advanceddffirentiation of the hlpostracum, which I
thicknesstowards the free edge as long as the mantle
should like to introduce here: In this class, a ring-
tissueis limited to the marginalparts of the shell. If the
shapedstructureuniting the shell epithelium with the
mantle fold covers the shell completely, however, the
shell,calledIhe "annulus",is associated with the head-
periostracummust show the samefeatureas the mother
foot retractors(Fig. 2a). This is the site of a special
of pearl layer, since the apex will have received a
secretion of "annulus material", a modified forn-rof
greater number of shell layers than the parts close to
mother of pearl. After removal of the animal, this
the margin (belemnites!).
material is recognizableas a dark ring of soft, poorly
The attachment of the soft body inside the shell is achieved (14) by the shell muscles or head-foot
calcified,conchiolin.It slowly progresses across(15) the earlier-formednormal layers of the hypostracum,
t/.
thus separatingthem from the posteriorones, which
definitive contents.(ln Sepia, where the septaare very
appear embedded in a special lining of annulus
closely spaced,the situation is different, however; the
substance.These posterior parts of the hypostracum
soft conchiolin first completely fills the future
will be called the "phragmocone",while the rest of the
chamber).Based on this observation,Appellof puts
shell (excludingthe periostracum)will be called the
forward the hypothesisthat in Nautilus and other forms
"conotheca".
Now, first we have to view more
with a typical phragmocone,the gas chamberswhen
closelythe relationshipof the annulussubstance with
first formed contain very "soft", i.e. very watery
the rnostposteriorparls of the shell: when the secretion
conchiolin. This hypothesiscan now be consideredas
of hypostracummaterial started,the annulusmust have
refuted. Indeed, the supportingridges, the pillar layer
been inserted directly on the primary, still
and the above-mentionedcuticula together form a
undifferentiatedembryonicshell, so that here it appears
morphological(genetical)unity, which is distinct from
to be connected to the ostracum(cf. Figs 2,14,16).
the septumand neck, on the one hand, and from the gas
The phragmoconeis not made of regular,tightly
space,on the other. The septalnecks show yet another
packed layers of mother of pearl. The latter are in fact
differentiation:the anteriorpart next to the septumis as
dissociatedfrom the ostracumand betweenthemselves,
solidly calcified as the septum,whereasthe posterior
so that gas-filled spaces,the so-called"gas chambers"
part in contact with the pillars is uncalcified or only
are formed. In one place the layers remain connected with one another,however, forrning a narrow tube, the
poorly calcified Iconnectingring]. Probably the "calcareouscones" are rather poorly permeableto gas,
"siphtrncttlar tube", which looks as if it pierces the
which can easily be transportedosmoticallythrough
"septa" of the gas chambers.Those parts of the
the "conchiolincones".This is indeednecessarysince
hypostracum forming the siphuncular tube are called
under the high water pressureto which the outer wall
"cones" fseptal necks] and are continuationsof the septa ". These septal necks do not really touch each
of the shell is exposed,some gas is extractedfrom the
another,sincethey are separated(much as the septaare
the accessopene.
chambersand has to be replaced.The pillars thus keep
separatedby gas) by a soft or loose mass with
Whereasin lhe endoceratidsthe gas chambersare
transverse"pillars". Where the septa insert on the
formed only dorsally and laterally (Fig. 2a) and the
annulussubstance,the open gas spacealso disappears,
siphuncle still completely fills the initial part of the
being filled by distally broadeningmother of pearl-like
shell, the other cephalopodsshow a much greater
shell layers,on which the septumseemsto rest. These
departurefrom the typical molluscan shell, in that the
shell layers are the "supporting ridges", which
initial part of the siphunclefcaecum]is connectedto
apparentlyare formed from secretionswhen in other
the shell wall only by a strengthenedpillar, the
parts the chamber gas is produced instead of shell
"prosiphon", whereas the rest appearsdetached(Fig.
substance.Pillars, gas and supporting ridges could thereforebe considered,as was suggestedby Appellof
2b) fNote: a prosiphon is uncertainin Orthoceras and absentfrom noutiloidsin general].
(1893),to be a morphologicalunit ("cavernouslayer")
The generalrole of the phragmoconeis clearly that
distinct from the septa with their septal necks ("main
of a hydrostatic apparalrls so far as this function
layer").
remainsunexploredin its details.But even after closer
This view is hardly coffect, however. It is true that
inspectionconsiderabledifficulties remain despiteour
in newly formed or incomplete (very thin) septa in
present state of knowledge. The great majority of
Ir{autilusthe pillars and supporting ridges are indeed
earlierauthors,e.g.Voltz 1836,Buckland1835,p. 631,
connectedby soft layers of very poorly calcified
assumethat (17) the gas content of a chamberedshell
conchiolin. Later these layers become (osmotically)
may changerapidly, since they generallyobservedthe
dehydrated,to finally be dried out to form the thin
septalnecksto be open to the gas chambers.Even Abel
"chalky pellicle" which covers the outer side of the
(1916,p. 168)contendsthat animalsmay rise or sink in
septalnecksand septa;we shall call it a "cuticula".
the water due to influx or efflux of water: others
(16) The gas-Jilled space has long been present,
believe that the gas is compresseddue to swelling of
however,so it must have been formed much earlier,on
the siphunclebetr"'eenthe septalnecks(due to injection
its own, and from the beginning it must have its
o f b l o o d ) . A 1 1 s L r c hp h y s i o l o g i c a l c o n t r o l s a r e
1 a I J
inconceivablegiven the real structureof the siphuncle
subclasswill be treatedin future monographs;here
(cf. Spirula, Fig. 28 andNautilu,s,Fig. 3a). A simple
only the following groups are presentedthrough
apparatusfor rising or sinking is not at the disposalofa cephalopod;the animal is thereforegenerallyconfined to a certain depth of water, so far as the phragmocone
their fossil representatives). 2. Subclass: DibranchiataOwen 1836. I.
determineshydrostatics.(The phenomenathat can be
O r d e r :D e c a p o d a L e a c h 1 8 1 8 o r t e n - a r m e d dibranchiates.
supposedto characterizethe formation of new gas
S u b o r d e ra ) B e l e m n o i d e a N a e 1 f 912 lthe
chamberswill be describedlater. when Nautilus will be
tetm occursat leastas early as 1894] or
discussed).
belemnite-likedibranchiates. Suborderb) TeuthoideaNaef 19 I 6 or squidlike dibranchiates.
C. Diagnosisof the Class
Suborder c) Sepioidea Naef 1916 or cuttlefish-like dibranchiates
The ancient cephalopodsare clearly characterizedby the above-mentionedpeculiaritiesof the shell. But there are younger types among them (since the Lower Lias), in which this typical apparatushas become largely degenerate, so we must look for other charactersto give a generaldiagnosis,which must then be basedon the soft parts. Sincewe know the soft parls only from Recent and closely related groups, the
il.
Order: Octopoda Leach 1818or eight-armed dibranchiates. Subordera) Palaeoctopoda Naef 1921 or palaeoctopus-like dibranchiates. Suborderb) CirroteuthoideaBer:ry 1920 or cirroteuthis-like dibranchiates. Suborder c) PolypodoideaNaef 1921 or octopus-likedibranchiates.
resulting insights are not necessarilyvalid for the whole class, so that a comprehensivediagnosisof the
In this arrangementwe conserveas many traditional
whole class is actually impossible. Therefore we
groups and names as is compatible with the
emphasize a characterization derived from the
requirements of a really natural system. For system
primitive form of the shell (Fig. 2). For the soft parts, as far as can be judged, the following statementscan be
meansorder and requiresa certain conservatism.As
made:
"Tetrabranchiata"are not recognizedas truly unnatural,
Cephalopodsare Molluscs, with one or severalarm crowns surroundingthe mouth, - with two mandibles
they must be conservedwith their names. (19) A
which togetherresemblean inversebeak of a parrot,
suborders;their names and definitions have changed,
with large catt€ta-€,v-€swhich are surrounded or envelopedby the basal parts of certain arms, - (18)
and they will be examined and discussed in the
with a "funnel apparatus"restricting the mantle slit,
Iong as the most generalized,useful groupings like
different matter is the traditional composition of the
following sections. We follow (also in the ensuingtext) the principle of
composed of a dorsal "nuchal attachment",lateral
naming families and subfamiliesafter "typical" genera;
"funnel pouches" and a ventral "funnel tube" made of
for higher groups the laws of nomenclatureallow total
two muscularlobesbent in or marginally fused to form a cone-liketube, - with large,yolky eggsundergoinga
freedom.But we neverlhelessrecommendthat even the
virtually symmetricalcleavageand developingdirectly into a young animal without a trochophore-llke larva.
higher groups be named after certain representatives, which once and for all should determine the significanceof the naming. When a group is split up, the name remains with the subgroup from which it originated, thus respectingnomenclaturaltradition. If
D. A survey of cephalopodsystematics
namesare basedon characters,all sorts of nonsense result once knowledge improves. For example, for
For the systematic subdivision of the Class
most of the "Tetrabranchiata" we do not know how
C e p h a l o p o dtah.ef o l l o w i n go v e r r r i e w isgiven:
many gills were present, and we have found "Myopsida" (cf. p. 30) having oegopsideyes(Spirula)
LSubclassT : e t r a b r a n c h i a t a : O w e n 1 8 3 6( T h i s
and "Lioglossa"(Ceph.Vol. I, Chapter48) with well
t4
Sch
Sw
J n.t. Ext.
Lm --Mg
--Bs '-0v
r:bl - "' Ps
Cs
Hz
Mr
Mt
Kr..Nk
Km N'd S s S Sw
'Ni Bs
I
TK T;
Fig. 3.
TK Mm
Tb
L
Diagrammatic cut-away views of Nautiltrs (a) and Sepia (b) illustrating the difference betwccn tetrabranchiateand
dibranchiateorganisation.Note: / . the lack of a ventral u''all of the living chamberin D and its replacementby the muscularmantle (Mrz), which conespondsto the insignificantmusculatureof the free cdge of the integumentalmantle (Mr);2. the envelopment(in b) ofthe shell by the shell fold, which coversthe outsideofthe shell with a secondaryshel1epithelium (Ss),so that the whole structure is enslosedin a "shell sac"; 3. the shift ofthe roofofthe mantle cavity in b, so that the anal papilla, the gills and renal sacs(etc.) are facing down- and forwards; 4. the loss of the upper gills in D, and the translbrmationof the brachial apparatus,the chamberedshell, etc. E"rl. outer arm crown; Int. inner arm crown, representedby prehensile(Ar, Tk) and buccal arms ("buccal firnnel")(Bl) in Sepia; Cl. brachialcirri; C.s.sheathof cimrs. togethcrwith the cirus it fbms one arm Lm. inner lip, sunoundedby the outer lip; Ofr.upper mandible; Zg. tongue with radula; Sk. buccal mass; Org. subradularorgan; Pg. pedal ganglion; lrg. visceral ganglion: Gg. cerebral ganglion;Nft. nuchal/collarattachment.adhcringto the mantle (Mt); Kk. hood; lo. anterioraorta;,rr.dorsalmantle groovc; Scft.shell; Sl. fleshy siphuncleinside shell siphuncle;Sr.r'.shell septa:Lh. coelom; G/. genital ligament; Mg. stomach;B.r. caecum;Ol,. ovary; Gs. genital septum;Ps. pericardialseptum;Hz.hear|' Ed. intestine; l'd. fore-gut (forming a "crop" in a); Vc. vena cava; Nl. kidney; Os. osphradium;Nd nidamental gland; Kn. gill ,4,ft.integumentalrnantle;Mr. mantle margin; Ifr. funnel valve: fi'. funnel tube.
l5 developedradularteeth.
epithelial"sheli sac" composedof a primary (inner)
When single specimensstand for species,specres for genera,genera for families, systematistscailing
and secondary(outer) shell epithelium, in which the position of the ventral part of the primary mantle and
them the types of the correspondinggroups, this practical use does not representthe rnorphological
the correspondingpart of the wall of the living chamberare occupiedby a thick muscularplate, called
norm in a strict sense;it merely servesto stabilize nomenclature.It would thereforebe betterto call them
the "mttscular mantle", - in which only 8-10 strong prehensilear[1s surroundthe rnouth and bear ,suckers,
"nominal types". But the usuai practice is apparently
at leastduring thejuvenile stage,- in which the eyeball
compatible with morphological principles and demonstratesthe superiorityof standardizationover
forms a closedcameraequippedwith an lrls fold and a lens, the whole embeddedin an orbital cavity, the free
mererecordinsof the known.
edge of which may (ontogenetically)contractto cover the eye either incompletely to form the "primary lid", or completely to form the "cornea",
E. Contrast betweenTetra- and Dibranchiata. Although we here conservethe subclassescreatedby Owen [1832], we have to emphasizethat only the subclassDibranchiatais basedon the recognitionof particular relationshipsof form, i.e. on a uniform, specialtype. Strictly speaking,lumping the other forms together in a SubclassTetrabranchiatacan only be justified in a negative sense,in that the characteristic
in which only one pair of gills is formed along with a distinct, dorsally situated"branchial spleen" which is attached along the greaterpart of its length to the muscular mantle by meansof a "branchial ligament", - in which the./unnel tube is lirmly closed along the ventral midline by fusion of the two halves, and interiorly partly lined with glandular cushions (the "funnel gland"), - the skin of which contains typical chromatophores(yellow, yellow-red, red, and brown)
featuresof the Dibranchiatado not occur in them. Types such as Endoceras, Orthoceras, Actinoceras,
permitting a rapid colour change(22), - in which an ink sac with an ink gland is present (at least as a
C y r t o c e r a s ,P h r a g n t oc e r a s , A s c o c e r as , L i t u i t es ,
rudiment) forming a rectal appendix.
Gyroceras,Nautilus and the Ammonoidea cannot be consideredclose relatives;they reflect a systematic diversityyet to be analysed.
In our opinion, the muscular mantle is an essential character of the Dibranchiata. Compared to the
But theseforms have a well preservedshell, i.e. an external shell with a typical subdivisioninto chambers,
Tetrabranchiata,its occurrencereflects a shift of the relationship between the animal and its shell. Substitutinga new, muscularorgan, greatly enhancing
(20) and thus they approachrather closely the general
mobility, for a protective shell meansan extraordinary
type of the class.The namesof subclasses(Ectocochlia
increase of effective range, opening up a number of possibilities for further modifications, as will be
and Endocochlia) suggestedby E. Schwarz (1S94) indeed make more sense.It would have been even better (21) if from the beginning they had been named after typical representatives(Nautilus and Sepia). But
demonstratedfor the different subgroups., The ot,igin of the muscular mantle is clearly expressed in
for the presentwe conservethe generally accepted
development:in embryonicdevelopmentthe initial site of its fixation (Figs 10,60) is alwaysthe free edgeof
names.
the shell (or shell sac).Subsequentlyvarious shifts may
In the following sections, we will be dealing exclusively with the Dibranchiata, which are clearly
occur(Fig. 7).
relatedto the older Tetrabranchiata as shownabove.
as a product of hvpodermal differentiationat the edge
The muscular mantle can thereforebe consideredl0 of the primary mantle (i.e. the typical, dermal mantle of the Tetrabranchiata)(Fig. 3). It reflects a thorough
Diagnosisof the dibranchiatecephalopods:
modification, and increasein effectiveness,of the locomotory apparatus:its contractionsrhythmically
Dibranchiatesare cephalopodsin which the shells or shell rudiments lie inside the body, ontogenetically
and intermittently reduce or expandthe mantle cavity, by alternatecontractionsof the circular and the radial
covered by a "shell fold' and thus enclosedin an
muscle fibre systems.Thus, water is sucked in and
t6 expelled, allowing at least respiration to continue
occurredin relation to Ihe envelopmentof the shell by
during periods of rest. Strongeractivity gives these
soft tissue.The shell fold and the primary (dermal)
pulsations a locomotor effect by means of the funnel
mantle are indeed connectedto one another at the
appararus.
apertureof the shell. But it is of course impossibleto
The funnel apparatusindeed occupiesthe entire
know at what rate (21) fhe two processeswent along in
mantle opening, narrowing it in a very peculiar way.
parallel, since there are no intermediateforms. Due to
Typically this apparatusconsistsof 4 parts: 1) the
the progressivedevelopmentand final closure of the
dorsal ntrchal attachment,which adheresto a smooth
shell fold at the end of the'conical shell, the latter
surface belonging to the primary mantle, called the
becamefully enclosedin a sac formed by the primary
"collar attachment" (Fig 3b), and thus fonns a tight
and secondaryshell epithelia(p. l3), a processwhich
closure,2) the ventral funnel tube, throughwhich water
also occursduring embryonicdevelopmentin all recent
is expelledwhen the mantle cavity narrows,3 [and 4]) the two l,aleralJunnelpouches,which then are inflated
dibranchiates.However, in the latter case the "shell
(23) and close the mantle slit-like valves. When the
outer shell never develops;the shell is alwavs formed
mantle cavity widens they come to lie againstthe body
insidethe sac(Fig.60).
sac" is closed before the shell is formed, so that an
and let the water enter quickly. The rhythmic shrinking
Due to their envelopmentby tissue the remaining
of the mantle cavity is achievedby different factors: 1)
parls of the shell are integratedinto the effective range
contraction of the inflated funnel pouches and of the
of lhe chromatophoreapparatus.The latter, along with
posteriorpart ofthe funnel tube, which narrow starting
the ink sac, constitutesa typical, specific, and indeed
from the front, 2) retraction,by rneansof the powerful
dynamicprotectivedeviceof the Dibranchiata.
shell muscles or head-foot retractors,of the entire
Given their normal position and architecture,the
head-footinto the mantle sacrL(Figs la and 2a), 3)
fins could also be considereda product of the muscular
contractionof the muscularmantle. Since thesefactors
mantle. Developmentalhistory, however, shows that
act rhythmically together in the Dibranchiata, a very
they must be consideredas hypodermaldifferentiations
considerableeffect is achieved:some forms move with
in the zoneof the shell fold (seep.32 andFig. 7).
such a power that they jet out of the water to cover
The muscular mantle, internal shell, fins, ink sac,
many fathoms ("flying squid"), very like flying fishes.
chromatophores,suckers,and completefunnel provide
In the Tetrabranchiata, there can be no muscularmantle
the animal with a mode of life, which
along with the
activity. Only the large funnel apparatusand powerful
highly evolved camera-like eyes
allowed the
retractorsare active and the effect must therefore be
Dibranchiatato be the sole invertebratesto openly
incomplete.But even in dibranchiatesthe functioning
compete with the vertebrates.They turn a chantbered
of this apparatusmay be simplified, e.g. by connecting
snail ntlo an ink/ish [Ger. Tintenfisch]!We will have to
the mantle more solidly with the funnel apparatus.
follow here the further metamorphosis of this
Along with this an increaseof mantle thicknessand a
organisationaltype. It involves discarding passive
descentof the funnel tube towards the body axis may
defencesand developingactive weaponsin the struggle
occur, so that the motive power becomes aligned
for existence.
(centered) with
We have no direct knowledge of the precursorsof the
the axis of
locomotion (cf.
Cephalopoda, Vol. I, Chapter32). Special muscles allow the funnel to be turned in
Dibranchiata.They should be expectedamongstthe orlhoceratids,whose phragmoconeclearly corresponds
different directions,so that a backstrokeof the ejected
with that of the oldest belemnoids. Evidence of
water resultsin movementin the oppositedirection.
developmentin the direction of the dibranchiatescan
Thanks to this arrangementthe animal can also swim
be seen in a reduction of the ventral shell wall
forward and turn sideways.For finer movementsthe
combined with a ventral position of the siphuncleand
fins participate, in particular as elevators, as do the
traces of a periostracum on the shell surface, as
united arms, some of which have special swimming-
observedin certain orthoceratids.The first and second
membranes.
featureshave beenjustly emphasizedby G. Steinmann
The evolution of the muscular mantle in the
(1910,p. 120) who found them in Orthoceras
ancestor of the Dibranchiata(Fig. 10a) must have
pletrrotomum Barrandefrom the Upper Silurian of
l7 octopodsare very poorly known from the fossil record. In contrast,the Decapodainclude numeroussignificant fossil representatives that must be treatedhere. First of all a generaldiagnosis: Decapods aredibranchiates inwhich l0 prehensile arms are present, the fourth pair of which (counting from the dorsal side) is more or less thoroughly modifred ("tentacular arms"); this modification is due to a lengtheningof the basal,sucker-lesspart into a "tentactrlarshaJi" andbroadeningof the distal, suckerbearing part into a "tentacular club", - in which the orifices of Ihe suckers (which are arrangedin two or Fig. 4. - An illustration of the difference between octopods and decapods.- Schematicrcpresentationof three juvenile
more rows) are surroundedby a stiffening "horny ring", (26) often with denticlessurroundingthe orifice
stages
(the median tooth can be transformedinto a hook
represcnting typical
p r o t o d i b r a n c h i a t e ,b )
arrangements: a)
decapod, c)
octopod. The
during post-embryonicdevelopment), in which a
protodibranchiate had five pairs of similar arms.the decapods
normal suckeris separatedfrom its muscular"suppoft"
have more or lcss markedly differentiated tentacular arms,
or "basal cushion" by a deep constriction,so that the connectionfonns a thin "stalk", - in which a distinct
the octopodshave only the four undifferentiatedventral pairs of arms. In no way do the octopodslack the tentaculararmsl (cf. the platesin vol. 2 ofCephalopoda.and below. p. 27).
crown of 6 to 8 small buccal arm rudiments ("buccal
dibranchiatcsand thus pcrmit considerationof a connection
pillars") connected to each other by a skin fold ("buccal membrane")is present("bttccalfunnel'), in
wtth Orthoceras.Adult octopodsat best show a plate-shaped
which the renal orifice.sare rrlore or less far removed
shell rudiment stiffening the dorsal part of the posterior
from the baseof the gills to lie closer to the anus, in
mantle. There are no distinct remainsof a phragmocone.The
which (except for the cranchiids) the funnel tube
embryo shows no more than the anlage for a rudimentary
containsa lunnel valye and connectswith the mantle on
shell ofthis type.
either side via an elongate,partly cartilaginous,scoop-
The decapods show many of the typical shell f-eaturesof
shapedadhesivedisk (."funnel attachment" or "Jilnnel cartilage"), similar in form and function to the nuchal Bohemia.But it seemsimpossibleto deducedescent lrom any parricularspecies. (25) A morphological element of dibranchiate
attachment(p. 23). This diagnosisshould be comparedwith the diagnosisof the octopodsin Part 5. Its mere length
organisationthat can be preservedin the fossil record is
indicatesthe comprehensivedifference and the strong
the beak. Such structureshave indeedbeendescribedin
uniformity of the secondorder.
considerable numbers and have been interpreted as
characterslisted here would not be recognizablein
parts of belemnites.All the "rhyncholites" that I have
fossil forms, even well-preservedones. and for this
seen (cf. Till 1907, 1909), however, belong to Tetrabranchiata(see Vol. I of Cephalopoda,Plates l7
reason some people have prematurely chosento base
and 18, and Fig. 42 below.,for the morphology of
on the number of arms.This was indeedan unfortunate
dibranchiatebeaks).
choice:in most fossil specieswe do not know anything
Most of the positive
the distinctionbetweendecapodsand octopodssolely
about the arms, and in the other speciesarms are much too incompletely preserved to allow any secure F. Contrast betweenOctopods and Decapods.
identification. In contrast,the typical structureof the adhesive
Within the Dibranchiata,we distinguish the orders
organs of the arms, which differs greatly betweenthe
Octopodaand Decapodaaccordingto Leach (1818).
two orders (Fig. 5), is essential and usefll for a
These very clearly distinct, natural groups are of
distinction of the fossil forms of octopods and
different importance for our study, because the
decapods.Certainly suckersare very rarely preserved
18 Niemiec (1885), which is now confirmed in its gencral
-st
aspects.
--*rr CL
--sk
in fossil dibranchiates(Fig. 85 [?62]). But the opposite is true for the products of transformation,namely the hooks,which are widely distributed,and it is important
sp
e so .
to note: 1) that recenthooks only occur in decapods,2) that the typical morphology of the decapodansuckeris theprereqttislle for hook formation. We can conclude from this that the fossil cephalopodsbearing hooks were decapods,even if the number of arms cannot be securelydetermined,or (27) appearsto be only eight or
_sk iu. y.
six. For in extant decapodsforms are also known that have only eight arms as adults, only six as early juveniles.The lattercaseoccursin all the "oegopsids", the fonner in the family Octopodoteuthidaeand in the
99' sp
genera Leachia, Chaunoteuthis and others. Intermediatestagesof post-embryonicdeveloptrent in decapodsalways show ten arms, however, the fourth pair of which (countedfrom the dorsal side) forms the
Fig. 5.
Typical octopod (a) and decapod (b) suckers.
Schematiclongitudinalsectionsthroughpart of an arm with a
typical "tentacular arms". These can be lost subsequently.
sucker. The latter is a special differentiation of the
But this is certainly not the case in the true
subcuticularconnectivetissue, the musculaturcof which is
octopods.They lack one of the dorsal arm pairs (cf.
linked to the muscular axis (ax) of the arm, whence (central ncrvous strand nr,) it receives a nerve which fbrms a ganglionic swelling (gg) below the suckerchamber(st). The inner lining of the sucker shows a "marginal ring" (rr), an
N a e f 1 9 2 1 , C e p h a l o p o d aV , ol. II, Pls 6 and 29), whereasthe arms correspondingto the tentaculararms of decapodsare presentas normal latero-ventralarms.
"adhesivering" (/rr), "wall-ring" (wr) and basalpart 1sp).The
Likewise these arms differ only slightly, or not at all,
latter fbms a muscular "sucking cushion", and the adhesive
from the other arms in the early juvenile stages of
ring is likewise equippedwith a strong muscular wall (hm).
decapods(28) (loc. cit. Pl. 23, Figs I and 2), and the
The n'all-ring is muscular only in octopods,whereas in
samecan be said of cefiain fossil forms (seebelow Fig.
decapodsit is stiffenedby a tough modification of the cuticle
el).
(.r) canied by a simple epithclium. In the octopodsthe whole lining of the sucking chamber(x) is a delicatecuticula which is regularly moulted. In decapodsthe adhesivering also carries a strong cuticula, which is organised(as in octopod
When comparingthe typical morphologyof suckers in octopods and decapods(Fig. 5), one inevitably realisesthat octopod suckersare simpler in structure,
suckers) into a system of platelets that can be moved in
i.e. less differentiated.A number of parts are lacking
relation to one another, since they are connectedby only a
(basal cushion, stalk, horny ring, denticulation)or do
very thin laycr. The sucker "support" fbrms a "stalk" and
not occur as distinct stluctures.Thus, they can be taken
"basal cushion" (bp) in decapods;its inside is made of a
to illustratethe primitive form from which the decapod
network of muscle fibrcs crossingone another,at the surface
suckersare derived,whereasderivation in the opposite
it shows an annularmuscle layer (rl), below which is a layer
direction doesnot make senseand would contradictthe
of longitudinal fibres (/g). Rg. annular musculature of the sucker,especiallydistinct in decapods;:. denticleson the free edge of the wall-ring; -t,.supportingridge; r,. depression in sucking cushion;ab. muscularbundlesallowing releaseof
o n t o g e n e t i c d e v e l o p m e n t .A p h y l o g e n e t i c i s tw i l l therefore consider the octopod suckers as the "primitive" form of decapod suckers (cf. Naef,
the sucker (by retraction of the marginal ring in the dorso-
Cephalopoda,vol. I, p. 98). The adhesiveorgans of
medialzone).
decapodsindeed appearmuch rnore advancedin terms
This rcpresentationmay be comparedwith the descriptionby
of complexity and efficiency: the differentiationof the
t9 supporting part permits free movement of the whole;
suckerson the three dorsal arm pairs, the two median
and when a prey animal attempts to free itself the
rows being transformedinto hooks. In these forms the
construction of the basal cushion and the stalk
ventral arms deveiopno hooks,the tentaclesonly a few
mechanically causesan increasein the sucking effect,
if any. If such an animal, e.g. Berryteuthismagister
since the stalk is pulling on the cushionwhich thus
(Berry) (Naef 192l, System) became fossilized,we
becomespafily extractedfrom the chamber.Likewise,
would recognizeonly three pairs of arms by double
in decapods,the mechanicalsupport of the sucking
rows of hooks, and we would thus obtain a picture like
chamberprovided by the horn,vrlng rendersmuscular
that proposedby Crick (1902, 1907)and Abel (1916)
action unnecessaryin this part. The denticulationof the
for belemnites.Fossil remains of belemnitesdo not
free rim acts like a series of claws when the sucker
permit any negative statement,however (cf. Crick
becomesattached,and the elevationsof the horny ring
1902,p.15).In most cases,they do not evenexhibitsix
prevent it from slipping. The horny ring, the denticles
distinct double rows of hooks. According to Huxley
and the knobs appearas specialthickenedstructuresof
( I 864) theremay occasionallybe 6-7 rows.
the delicatecuticula that is also presentin octopods.
If Acanthoteuthisspeciosa(Fig. 9l) really tums out
The mechanicalpropertiescan be seenparlicularly well
to be a belemnite,as was supposedby Miinster(1830)
when suckersare studied in dead animals. Even in decayinganimals- 24 hourspost mortem- the suckers
and strongly supportedby Angermann (1902), there can be little doubt that ten arms were present in this
still function when toucheddue to mucousadhesionof
animal. The same can be said for Belemnotetrthis and
the marginal ring, and when one pulls on the adhering
Phragmoteuthis. As far as can be judged today, the
sucker,the sucking effect increaseswith the changeof
fossil belemnoidshad ten similar arms, each with two
partial pressureof air and water, due to the resistance
rows of hooks. It is not yet clear whether the soft parts
of the stalk and the withdrawal of the cushion.Similar
already showed signs of tentacle differentiation,
effects occur only very incompletelyin octopods,so
something which I previously assumed (Naef,
that the functioning of their suckersis almost entirely
C e p h a l o p o d a v, o l . I , p . 1 1 0 , 1 3 0 ) . B u t t h e t o t a l
d e p e n d a n to n m u s c u l a r a c t i v i t y ( c f . N a e f 1 9 2 1 ,
correspondencein this respect between the mutually
Cephalopoda, vol. I, p. 121),and it ceasesafter death.
independentRecent (derived) types (Teuthoidea and
As indicatedabove,the denticlesof the homy ring
Sepioidea) indeed suggestsit. In any event, (30)
act like claws, their effect being more or less markedly
differentiatedtentaculararms must be assumedfor the
enhancedby their form. This is achieved,for exarrple,
common original forms of the recent decapods (Aulacoceratidae?).
by enlargement of the distal median tooth (cf. Cephalopoda,vol. I, Pl. 12, Fig. 8). (29) This tooth, when growing strongly during postembryonic development(loc. cit. p. 131) may finally becomea powerful hook that cancels and replacesthe sucking
G. The subordersofthe Decapoda.
effect ofthe original sucker.This developmenttendsto
I have subdivided the decapodsinto three natural,
involve only part of the suckers and arms, thus
clearly distinct suborders(Naef 1921, System),largely
resultingin a division of labour.The remainingsuckers
following earlier systematistsand using a terminology
indeedtend to form no denticlesat all and thus lose anv
which had been usedearlierwith differing, often vague
claw-likefunction(loc. cit. p. 130-132).
meanings; these suborders embrace both recent and
When in fossil cephalopodsdouble rows of hooks
fossil forms. In doing so I have opposedthe dominant
appearinsteadof arms (Fig. 68), one can be certainthat
practiceof treating fossil forms separately(Zittel) and
a decapodis under consideration,but one does not
adopting the subordersof d'Orbigny (1845) for the
know how many arms were present in the animal or
Recentforms. The latter author presentedthe decapods
how many rows of suckerswere originally presenton
as subdividedinto Myopsida and Oegopsida,a blunder
each arm. Tracesof more delicatesuckersare virtually
which indeed generateda great deal of unprofitable
never well preserved!When trying to reconstructthe
discussion.Its rebuttal, with all the necessary
animai we have to rememberthe conditions in certain typical oegopsids(Gonatidae),which have four rows of
arguments,is presentedin volume I, chapter 5 of Cephalopoda. In the present work the artificial
20 forms under belemnite-like forms, or Belemnoidea, with the following diagnosis:belemnoidsare fossil decapods having a well
developed, straight
phragmoconeat the end of the mantle sac; having (as far as is known) hooks insteadofsuckers on all (10) or at least on some arms. (The latter limitation might become obsolete for some forms in the event of new For the time being it remainsvalid). observations. The typical shell form of belemnoids,forming a is shown in Fig. 62. In solid basisfor our systematics, this structure,which is entirely internal (as is the case in all the dibranchiates),we distinguishthe typical layers of a molluscan shell (ostracum, hypostracum, periostracum) and the typical chamber formation of cephalopods.The periostracum is representedby the "sheath",which tapers into the terminally thickened "rostrum". The hypostracumforms the inner stiffening layers made of mother of pearl; the septa (3 1) and siphuncular funnels Iseptal necks] appear as differentiationsof this stiffening layer. The ostracum, together with the adjacentlayers of the hypostracum, surrounds the chambered "phragmocone" and forms the so-called"conotheca"which continuesanteriorlyas a tongue-shapes"pro-ostracum".The iatter can be 5
Fig. 6.
Schematicrepresentationsof teuthoid and sepioid
shells,illustratingtheir morphologicaldifferences. Prototypes of
a)
Decapoda, b)
Teuthoidea, c)
consideredas the dorsallimit of the living chamber,the ventral limit having been replacedby the "muscular mantle" (Cephalopoda,vol. I, chapter2, p. 92-94). The
transitionalform to sepioids;/)ancestralform ofSepioidea.
whole ostracumtypically grows by marginal accretion and - accordingto the type of accretion- showsup as
Metateuthoidea;d) transitional form to belemnoids, e) Actual genera are: Nannobelas Pavl. (a), Paraplesioteuthis
the mid-dorsal plate, the lateral plates and the ventral
Naef (&), Palaeololigo Naef (c). Diploconus Zittel (d1.
plate. While the other parts grow rapidly by anterior
Belemnosellan. gen. (/).
accretion, growth is much slower laterally and
The mantie sac and she1lare shown; the she1lis shown in
especiallyventrally. Even when the pro-ostracumis
white, as if transparent,the muscularmantle is crosshatched. Note the degenerationof the phragmoconein the teuthoids (b, c), the penetrationofthe shell apeftureinto the mantle sac and the shift of the insertionof the muscularmantle on to the
missing,this processcan be clearly recognizedfrom the growth lines of the conotheca;the different zones of growth being demarcatedby rlore or less distinct
sheathin sepioids(e,l).
longitudinal lines. Reflecting the courseof the growth
-1.median asymptotes;2. lateral asymptotes,3. lateralplates;
lines, the rnedianplate is also called the "parabolar
4. ventral mantle insertion;5. dorsal mantle insertion(latera1
field", the lateralplate the "hyperbolarfield". The lines
margin); 6. free margin of sheil; 7. pro-ostracum.
limiting the latter are called the median and lateral "asymptotes",disregardingthe proper sense of the telIn.
distinction is replacedby a natural one also applicable
The insertionof the musctrlar mantle foilows the
to the fossil decapods.The latter are recognized as
free margin of the shell, as is the casein the errbryos of
forms having hooksinsteadof suckerson the arms.
all recentcoleoids(p. 22).Clearly the shell is merely
a) Among the extinct decapods,most speciesand
coveredby a skin fold; it is not "surroundedby the
individuals represent a type that is rather purely
mantle" (32) as generally stated. In the embryo this
expressedinthe belemnlles.We subsumeall the related
"shell fold" alreadyproducesmuscularstructuresin the
2l form of fins; thus they are originally situated on the
of this type (33) has not yet been recognized;it will be
outer surfaceof the shell, whencethey can later depart
of special interestfor our survey. Like the Teuthoidea
( p .3 6 ) .
it must be derived from the Belemnoidea,though in a
b) Some fossil decapods are related to recent
very different way: imagine the shell of a belemnoid
loliginids and have therelorebeen named"loliginites"
(Fig. 6d) becoming very bulky, so that a shift of the
by Quenstedt(1849).We now call them calamary-like c e p h a l o p o d so r T e u t h o i d e a ( T e u t h i s , T e u t h o s i n Aristotle's writing : calamary).They are distinguished
insertion of the muscularmantle from the free margin along with a ventral curvatureof the phragmoconethat
from the above-described type in particularby the total
is also recognizablein most belemnitesand in the
degenerationof the phragmocone,which no longer
poorly known Diploconus (Figs 65, 71), which might
of the shell to the outsideof the sheathis conceivable,
shows distinct septa,siphuncleor chambersand which
be an intermediateform. Both these modiflcations of
is also strongly reduced in size compared to the muscularmantle (.p.2a).For the rest, the same shell
the typical shell characterizethe fossil and recent
parts are still recognizable(conus and pro-ostracum,
modifications or reductions. They are further
medianand lateralplates,rostrum and asymptotes),and
emphasizedin an ideal intermediateform (Fig. 6e), in
the generalcharacterof these parts may still be so
order to illustrate the real, typical situation (Fig. 6f) of
strongly reminiscentof the belemnites(Fig. 6b) that
primitive Sepioidea(Belentnosella).Thus we seethe
s p e c i a l i s tl si k e V o l t z ( 1 8 3 5 ,1 8 4 0 )a n d A g a s s i z( 1 8 3 5 )
achievementof a rather peculiar shell lorm which is
Sepioidea,as far as there are no further (ensuing!)
declaredthem to be belemnite remains. In contrastto
characterizedby (1) the formation, on the surface of
these "Prototeuthoidea"(e.9. Geotettthis,Leptoteuthis,
the sheath,of ridges and edgesenforcing the insertion
Plesioteuthis)we find a closer relationshipto recent
of the muscularmantle, while (2) the shell opening, l.
forms in Trachyteuthis,Beloteuthis andPalaeololigo,
e. the free margin of the shell, migratesto the inside of
the last genus (Fig. 6c) demonstrating the most
the soft body, generating far-reaching anatomical
complete achievement.We therefore designatethese
modificationsthere (Spirula'.Naef 1913,p. 454-461.).
forms as Mesoteuthoidea,while the recent, more
Along with this process(3) a strong ventral curvature
advancedforms are called Metateuthoidea(Loliginidae
of the phragmoconeoccurs, which in any casewould
and allies, oegopsids).In all theseforms the mid-dorsal
prevent a belemnoid-type connection of
plate, which is already tapering anteriorly in the
phragmoconewith the body of the animal (Fig. 6a).
Mesoteuthoidea,becomesincreasinglynarrow while
This new type, when comparedwith the belemnoid
the lateral plates broaden;the conus becomeseven
type, also representsa decreaseof passive shell
more reduced (often flat, spoon-shaped)and the
apparatusagainst an increase of the active elements
broadenedlateral plates together with the remaining
(muscular mantle) (p. 22), furthermore stabilizing the
parts ol the conothecaform a foliate structure,which is
hydrostatic equilibrium, since the large gas chambers
called the "vane" in contrast to the median rib.
are situtatedin the dorsal part of the body. Among the
the
representing the mid-dorsalplate,which is now called
Sepioideathere are fossil generaltke Belentnosella,
the "rhachis". On either side the vane is subdivided.
Spirulirostra, Spirulirostrina and Belosepia, which
more or less distinctly, into the lateral plate and the "conus vane", the two being distinguishableby the
representthe types of as many families, forming a kind
persistent"lateral asymptote" (Cephalopoda,vol. I,
are Spinrla, Idiosepius and Sepiola. These forms are
chapter4, p. 146, Fig. 62). The recent shells of this
closely relatedto one anotherand, via the shell form of
type no longer show any calcification;the fossil shells
Spirula,linked to the fossil Sepioidea.
were at least partly calcified, and the prototeuthoid
of seriesleadingto Sepia. Recentgenerato be added
The total independenceof the subordersb and c is
shells were probably always calcified. The shell of
endorsedby palaeontogy:although the sepioid type is
Teuthoideais calledthe "gladius".
much younger(Eocene)than the teuthoidtype (Lias), it
c) Again few fossil but numerousliving decapods
conservesto the presentday the ancestralchambered
are closely related to the common cuttlefish; we
shell (phragmoconewith prosiphon,siphuncle,septa,
thereforecall thern Sepia-hkedecapodsor Sepioidea.
etc.), either without rnuch modification (Spirula), or
Up to no"v the morphologicaland systematiccharacter
strongly modified but well
d e v e l o p e d , ( 3 4)
22
Fl r
F[r t
Ss
F t "
Fle
; -l4i
b
23 Fig, 7, - The morphology of the relationshipbetweenthe shel1(Sc/r),the shell sac (Ss),the muscularmantle(Mm), the coelom (Co), gonad(Hd) and fins (F/) in differentdibranchiatet)?es, illustratedby crosssectionsthroughthc posteriorend of the body. a) Generalizedinitial condition in dibranchiates:thc fins with their smoothfin cartilage(Kr) rest directly on thc outer surfaceof the shell,nearthejunction of conothecaand pro-ostracum.(cf Naef, Cephalopoda, vol. I, p. 95). b) ldeal initial condition in decapods:as can be seenin embryos.the part of the primary shell sac that carriesthe fin cartilageor', cither side becomesseparatedby lblds which form the basal lin pouch (IIlt), so that the gliding arliculationbecomesindcpendentof the surfaceofthe shell. c) In recentforms this separationis complete,thus offering the option of moving the finbases on to otherparts of the shell (fl orthe muscularmantle(e). d) The latter state is found in the teuthoids (c-d), in which (in the course of subsequentdevelopment,generally during the postembryonicphase)the muscularmantle largely envelopesthe shell. leaving the primary insertion site behind at the shell margin (Part l, p. 22); thus it comes to lie between the fin pouch and the shell, which may ultimately be covered completely, or almost completelyfby the muscularmantle],leavingonly the mid-dorsalkeel free (K/). e) Total enclosureis achievedearly in the genusLoligo (Kt). so that the gladiuslies inside the cylinder of the muscularmantle,while the fins lie outsidcand aremobile using the specialintegumentalmusclesassociatedwith them. fl In the sepioids(especiallyin Spirulirosta) onc can assumethat the lateral margins of the shell (Sfl become suppoftsfor the fins (cf. Figs 8,11,22); from this conditionthe fins lengthenin the sepiidsto reachthe anteriorpart ofthe muscularmantle. Fa. vane of the gladius(p.32); Gl. genitalligamentlFl1 and Fl2,upper and lowcr opponentswithin the fin musculature.
(35/ containing gas and being fully calcified (Sepia).
systematists.The sliding is made possibleby a sort of
(The Teuthoideahad alreadylost the typical chambers
articulation;the inner face of the cartilage and the
in the Liassic period, i. e. when they first appearin the
substratumboth belong to a flat epithelialpouch which
fossil record).
will be called the fin-base pouch. This structure exhibitsvarying topographicrelationsin that it may lie on the shell or on the muscular mantle. The latter
H. Morphological relationships between the shell
situationseemsto be more comrlon and might suggest
and the fins.
that the fins were derived from the muscular mantle. The finer structureof the fins would indeed argue in
For the reconstructionof fossil decapodsthe fins are of
favour of this derivation. However, development
great significanceboth in terms of overall morphology
suggeststhe following relationship:
and ecologicalrelationships.I have seena numberof
The fin-basepouch is a separatedpart of the shell
fossils (Figs 42 and 52) which show the fins very
sac and its epithelium thus stems from the secondary
clearly and thus permit an interestingcomparisonwith
shell epithelium(p. 24). This is true also for the part
recenttypes. To understandthe insertion of the fins in
subsequently stiffened by the cartilage and thus
fossil decapods, one obtains some unexpected
becoming a sliding surface. ln general terms the
information from the developmentof living decapods.
"morphological place" of the fin rudiments is the
Here I
provide some preliminary
outside of the shell, and their material originally
observations,and I must refrain from giving a special
belongedto the connectivetissueof the shell fold. The
illustration(see forthcomingvol. lll of Cephalopoda,
fins are indeedtypical for the dibranchiatesand remaln
a n d v o l . I , 1 9 2 1 , P l a t e s2 - 5 , 1 2 , 1 5 , 2 3 , 3 7 ) . T h e
in closeconnectionwith the intemal shell (cf. Fig. 4).
can only
following observationsare pafticularly noteworthy:the
When trying to visualizethe primary morphological
f i n s o f d i b r a n c h i a t e s ,e s p e c i a l l y d e c a p o d s , a r e
characterand subsequentmodification of theseorgans,
proximally supportedby a cartilage. The latter slides
we have to take the following view: on the outside of
66) on a smooth substratumand allows a shift of the
the conical shell longitudinal extensionsappeared,
whole fin, effected by muscular contractions,so that
originally in the form of modest folds that were
the fins are not constantly kept in the same position;
supported by the musculature integrated in the
this holds true also for preservedspecimens a fact
connectivetissueof the hypodermis.In the courseof
that has been overlookedby earlier morphologistsand
their further differentiation as persistentstructures,
24 their connectionwith the substratumbecameboth more
A. Preliminary remarks.
intimate and more mobile. First the fln cartilageswere sliding on the shell. In the decapods,at leastlr, the
In beginning this study with the palaeontologically
associatedpart of the shell sac later became separate
youngestgroup,we have different aims in mind: first, a
and thus formed the fin-base pouch, which allowed a
clear presentationappearsespeciallynecessaryas their
further removal from the shell. This occumednot only
p e c u l i a r i t y " i s n o t s o e a s i l y u n d e r s t o o da n d h a s
in the teuthoids, where the muscular mantle
thereforebeen misjudged.Secondly,the group includes
secondarilycomes to lie betweenthe fin-base and the
some of the most .frequentlycited and indeed most
shell,but in the sepioidsas well (Fig. 7a-f).
accessibleforms of cephalopods,the morphological
The issue is of great importance for our
understanding of which is particularly desirable.
considerations, since a primary relationshipbetween
Thirdly, are we able to offer here many ne\Nelements
the shell and the fins is obvious (37) and can also be
and thus have an opportunity to demonstratethe
applied to fossil shells. (See for comparisonof details
successful application of some methodological
Figures7, I 1, 18,25, 39,42, 59, 61,62, 67 etc.).
principles(cf. above,p. 4).
The primary position of the fins in recent decapods is at the posterior end of the mantle sac; wherever a
Diagnosis.
conus is presentfins lie outsideit. This is essentialfor the reconstructionof fossil forms, especially since
Sepioideaare decapods, in which the phragmocone,
some of them (Figs 42 and 43) show a very similar
as far as it still exists, is posteriorly incurved ventrally
situationto recentjuvenile stages(Fig. 61). In both
and the free lower edgeofthe conothecais pushedinto
instancesthe fins are very small. It is only secondarily,
the insideof the body (Figs 9-11) so that the insertion
in most casesduring post-embryonicdevelopment,that
of the muscularmantlecomesto lie on the outsideof
they can grow forward on either side of the mantle to
the sheath, in which the posteriorfin edgesare not
approach a state sirnilar to that known in Sepia
united (Fig. 31),
(Cephalopoda, vol. I, Pl. I and 2). This most widely known form of decapod is a very derived type,
perforated,i.e. has no longitudinal canal between the afferent and efferentvessels(39) (Frg.36), - in which
however, which has long induced palaeontologiststo
the radularteetheachhavea singleblunt cusp.
erroneousideas about older types (belemnites).Much
in which the axis of the gills is not
When viewed in comparisonto living Teuthoidea,
of what has been interpretedas fins adjacentto fossil
the Sepioidea show numerous negative features of
mantle complexes are, in fact, extruded or decaying contentsolthe mantlesac(cf. 75-76\.
minor detail: well preservedshells that are calcified, absenceof neck folds, etc. (cf. Cephalopoda,vol. I, chapter38).
(38) Part II:
B. Introduction.
The Sepioidea or sepia-like Before giving a general characlerizationof the new
dibranchiates.
suborder,and a description of their representatives,I Contents'.A. Preliminary remarks and Diagnosis(p. 38). B.
should like briefly to illuminate the necessity of
Introduction.C. The typical organizationand developmentof
creating this suborderand show the way by which I
the Sepioidea@. a3). D. The family Belemnosidae(p. a8). E.
have come to this conclusion.
T h e f a m i l y B e l o p t e r i d a e( p . 5 3 ) . F .
The family
Belosepiellidae(p. 60). G. The family Spirulirostridae (p. 60). H. The family Spirulidae(p. 68). I. The family Sepiidae (p. 79). L. A review of the fossil Sepioidea and their evolution(p. 94).
This is of some
interest,sinceit revealsthe leadingideasof systematicmorphological considerationsand their application to palaeontology. The nomination of the sepioid type is a surprising result of systematic-morphological analysisbecausethe respectiveforms have long been known; they belong to the most frequently cited and most thoroughly studied
25 closely related fossils (,Spirulirostra, Beloptera) was completelyoverlooked(cf. Lang 1900,p. 95, Fig. 107). In contrastto this situation,it must be mentionedthat the classicalmalacologists,e.g. Blainville (1825) and d ' O r b i g n y ( 1 8 3 9 , 1 8 4 1 )r e c o g n i z e dt h e e x i s t e n c eo f theserelationships(cf. Cephalopoda,vol. I, chapter38, on Beloptera and Spirulirostra). Incidentally, I began my own studies under the burden of some further erroneous assumptions.The most unfortunate of these was the firm grouping of sepioid types and loliginids as "Myopsida", placed in opposition to the "Oegopsida". Only after moving beyondd'Orbigny's (1845)mistakein this respect,the right way forward for a linkage was found: the study of the embryonic developmentrevealedthat the iuvenile shell of Sepia (Cephalopoda, vol. II, Pl. 20) is much more closely relatedto the recent genusSpirula than to the fossil belemnoids(Fig. 8a, d). A strong similarity lies in
the strongly excentric shell growth,
characterizedby fastergrowth on the dorsal side.Ifthe Fig. 8.
Morphological relationshipsbetwecn Spitula and
Sepla illustrated by diagrammatic figures of four juvenile stages from the genera Spirtrla (a), Spirulirostra (b), Belosepio (c) and Sepicr({. a is a planktonic young animal,
shell curvatureis strong from the outset (protoconch), as in Spirula, ventral coiling results. If it is feeble (Sepia), ventral incurving of the shell virtually
drawn from nature; c is a corresponding,advancedembryo (cf. Cephalopoda,vo1.II, Pl. 19); b and c are reconstructtons
disappears.In both instances, however, the free, primary ventral edge of the shell is directed towards
from the juvenile stagesthat can be recognizedin the fossil
the inner part of the mantle sac.Due to its flatnessand
shells. For a morphological interpretation begin u,ith b in which the pro-ostracumis presentin its typical form.
to the belated differentiation of the rostrum, the "hump". the "fork", and the "dorsal plate", the cuttlebone(cf. p. 86) looks so different from the shell
stippledareas,but the [integumental]shel1fold is assumedto be transparent.as it is in very young Spirula and in Sepia
of Spirtrla that the latter appearedmore ciosely related
The muscular mantle and head-fbot are representedby the
embryos when the chromatophoresare conffacted, so that the shell can be seenthrough the skin in considerabledetail. (In all these juvenile forms the sheath forms only a very thin corer ol'the phragmocone ).
to the belemnoids, whereas the former would be c o n s i d e r e dm o r e c l o s e l y r e l a t e d t o L o l i g o . I t i s undeniable,however, that Spiru/a and Sepia have in coilrmon the chamber formation and calcification of the shell, so that a basic similarity exists.Furthermore, the anatomicalstudy of Spirula (cf. Cephalopoda,vol.
forms of the class (Sepia,Sepiola, Spirula). They will neverthelessappearas thoroughly misunderstood,due
I, and Chun 1910)revealeda very striking similarity with Sepia (41) and its closestrelatives(Sepiolidae),so
partly to inadequacy of methodology, partly to
that the systematic-motphological need for bridging the
objective difficulties.
gap in terms of shell formation as well became
In (40) particular the lack of
knowledge and consideration of fossil forms by zoologists,and of living forms by palaeontologists,
related animal,s could only be ttnderstood via
resultedin little understandingof generalrelationships - a circumstancethat must be again emphasizedhere,
this is really the case. This figure presents
as it representsa widespreaddrawback (p. 3). One
reconstructionsof young Belosepia and Spirulirostra,
should remember, for instance, that up to now (see
which were made by first drawing the isolatedjuvenile
Riefstahl 1886).Seplawas comparedto belemniteswith
shells (accordingto a and d) and then adding the soft
some lack of caution,whereasoften its affinity ro more
parts in their typical form. It appearsthat the juvenile
obvious. Indeed, the opposition of parts in so closely intermediaty stages.Figure 8b and c demonstrates that
26
Fig.9. - Schernaticmedian section through thejuvenile stageof Spirulo. drawn in 1913 from a preparationin the possessionof C.Chun(Leipzig).(ct'.alsoChun 1915,Fig. 1. Pl. 73). Note the strxctureand position of the shell inside the body. especiallythc orientationof the shell apcrturewhich is pushedinto the visceraand embracesthe liver, and note also the peculiar inserlionofthe muscularmantle ventrally and dorsally on the outer surface ofthe shell. Anothcr striking feature is the posterior lengtheningofthe dorsal mantle cavity which extcndsbeyond the end ofthe shell; a typical pro-ostracumis not only lacking, but is indeed incompatible with this arangement. The mere presenceof the muscularmantleat the mid-dorsalline is, of course.a remarkableabnomality tn a decapod.(cf. Fig. I 0). I-6. gaschambers;Ps. prosiphon;Sa. initial caecumofthe siphuncle;Sd. septalneck; Sw. septum;Sc. siphuncularcoelom; Sl. fleshy s i p h u n c l eF;g . f i l l i n g t i s s u e C ; o7-Co:.paftsofcoelom G;o . G o n a d . - r . v c n t r a l w a l l osf h e l l ;M a . s t o m a c hB; / . c a e c u m H ; z.heafi;Gg. gastricganglion;Nl. kidney; Mrz. muscularmantle;Ed. intestine;7b. ink sac;AJ. anus;Mv. mantle cavity; Vc. vena cava; Ib. liver; Ao. aorta;Md. dorsal mantle cavity; Oe. oesophagus;Gd. poison gland; St. statocyst;l/g. visceral ganglion;Nft. nuchal attachment; Cg. cerebralganglion;Pd. pedalganglion;06. upper buccal ganglionl Lb. lower buccal ganglion;Dr'. funnel gland; Ifr. funnel valve: Ir. funnel tube;l/. outer lip; ,Sr.subradularorgan;1/. inner lip; L'&.lou.crmandible;Ot. uppermandible;Rd radularsac.
stagesof thesetwo fossil forms fit in perfectlybetween the juvenile forms of Sepia andSpirula. The latter thus
the conservedventral cLtrvalurerelatedto this (p. 40)
appearas 0,pical voriants of a commonprimctrl'.fornt, the most essentialfeature of which is the ventral
margin into the viscera. An additional feature is the
curvature of the phragmocone.Apparently this forrn
subsequently(Fig. 32) becomesconical, whereasin
was fully achievedin Spirulirostra. Frorn this form Spirula may be derived by suppressionof the pro-
Sepia (Cephalopoda,vol. II, Pl. 19) it is still distinct in the juvenile part, but only a shallow depressionfrom
ostracumand subsequentpenetration(42) of the dorsal
the very beginning.
as well as the sharp penetration of the conotheca tubular initial part of the siphuncle, which only
shell margin (Fig. 27) into the soft body, with even
The significanceof thesedetailsbecameclear when
strongerventral curvatllreof the shell, which later leads
I looked at sagittal sectionsof Spirula (Fig. 9) and
to its spiral coiling. A degenerationofthe pro-ostracum
realized the particular relationshipthat such a shell
is also recognizable rn Sepia, but in a different
must haveto the soft body.
connection.The dorsal shell margin remains in its
A further comparisonof theseconditionswith those
original position; in contrast,the gas chambersare
of recent and fossil forms taught me that Spirula
progressively shifted forward until they reach the
characterizesa well defined group of decapodswhich
dorsal shell rim, so that no space remains for the
vary considerably,but neverthelessare very similar to
formation of a distinct pro-ostracum.In drawing a
Spirula in a number of featuresthat distinguish them
suggestionof a pro-ostracumin Fig. 8c, I merely make
from other decapods.Indeed,in addition to the above-
an assumption.Belosepiaalreadyis a sepiid'3,although
mentioned peculiaritiesof the shell there are some
i t m o r e s t r o n g l y e x p r e s s e st h e g e n e r a l s e p i o i d
independentcharacterswhich permit an unambiguous
character.
diagnosisof the Sepioidea.
(13) Thrs concerns the stronger curvature of the protoconch(which is no longer globular,however),and
27 The metamorphosis shown by these figures appears straightforward and easily understandable,because no periostracalelementsare present.Such clementsonly appear
a.
at later stages,but they must alreadybe taken into accountto understandall the changes,sincethey may partly have caused them, especiallythe stepleadingfrom c to d (cf. Fig. 6d, e).
b.
C. The typical organization and development of the Sepioidea. 5a
-_11
The recent sepioids - with the exception of the
c.
somewhatlarger Sepiidaeand Rossiinae are always
11
very small animals, measuring in general only a few centimetres in
length, that
live
(save for
Heteroteuthinaeand Spirula) mostly in the coastalzone (1-400 m depth), on the sea bottom, partly swimming, partly resting or even buried in the sand and mud so
d.
that only the eyes and the funnel openingsremain free. In this position they lurk for prey which is seizedby ejection of the tentacle arms. More commonly, 1 \
however,they move aroundwhen hunting, generallyat
Schematicdiagrams of the derivation of sepioid
night. This typical mode of life can also be assumed(as
organisation.Idealised mcdian scctions through four young
far as the shell forms can tell us) for the fossil
cephalopods(mantlesacs)with two gas chambers.
representativesof the group, which were very small
Fig. 10.
o) Orthoceras grade. External shell, wall of living chamber complete,wholly cnclosingthe mantle sac;roof of the mantle cavity facing forwards and upwards; anal papilla situated closeto the mantle(cf. Fig. 3a). 1. first septum;2. second septum;3. first septalneck; 4. secondseptalneck; 5. dorsalwall ofliving chamber;6. dorsal
decapods,as will be shown.They first appear(p. 33) in except Belosepia - remain scarce!
the Eocene and
although some of them were widely distributed in Cenozoicmarine sedimentsof the littoral zone.(44) The mode of developmerl is conditionedby the
part of mantle; 7. anal papilla; 8. ventral wall of living
relatively large eggs of the sepioids.The considerable
chamber; 9. ventral part of mantle; 10. shell margin; .r.
egg size resultsin a very completedevelopmentinside
mantle margin.
the egg case.The recent forms therefore tend to look
D) Hypothetical transitional stage. The free mantle margin
similar to the adult animalsalreadyat hatching(but see
has produceda musculardifferentiationventrally, the anlage
Spirttla, Fig. 27). The relationshipsbetween the shell
of the muscularmantle (12). and a distinct she11 fold (11)
and the mantle sac that are typical for Sepioideaare
beginsto cover the outer surfaceof the shell, which a11owsa betterconnectionof the muscularmantlewith the she1l. c) Dibranchiate(: decapod-belemnoid)stage.The muscular mantle (12) has largely replacedthc ventral wall of the living
establishedduring the embryonic phase (Figs 9 and 10). Moreover the typical organisationof adult sepioids
chamber and the associatedprimary or "integumental"
will be described.The aim is to combine in an overall
mantle, exceptfor a l-ewremnants(at l0a), and has taken the
picture the rnorphologically primary charactersthat
roof of the mantle cavity with it in a posterior direction,
have been recognizedin the courseof the comparative
whereas the anterior part with the anal papilla (7a) is shifted
analysis of existing diversity. The ideal form thus
forward.
construedis called Protosepioides,assumingthat it
fl Sepioid stage(Spirulirostra). The phragmocone is bent ventrally in the posterior end of the mantle, the free margin of the shell penetratingthe body, whereasthe muscular mantle (12) has shifted its insertion (13) on to the outer surfaceof the she11.
largely resemblesthe real ancestralspecies(seebelow, p. 49).We will describeit only briefly, referringthe readerto the presentFig. 11 and to the more detailed treatmentin volume I, chapter38 of "Cephalopoda".
S = = = -: '= ? - a= . . = 4? : t r ' l
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29 Protosepioides,i.e. the prototype(ideal form, type) of the Sepioidea,is a decapodwith an overall aspect similar to the living genusldiosepius.
wherecomelatedmodificationsare initiated(cf. p. 33). 4. The phragnloconeexhibits typical featuresas far as details are concerned,but shows a marked ventral
Major differencesof this prototypefrom sepiidsare
curvature especially in its earliest part, which is very
the small size and the terminal position of the rounded
characteristicfor the group and which is relatedto the
fins, and the rather narrow mantle sac (conditionedby the shell form) which is typical for sepioids.The shell
above-mentioned shift insidethe body.
correspondsperfectlywith the shell of Belemnosella
formation of a massiverostrlrnl,which housesthe early
americana (Fig. 12), as far as the latter is known, and
parts of the chamberedshell. The rostrum in its turn
should have had a delicate,flexible pro-ostracum. The
could be relatedto the benthic life style, which requires
phragmoconewas probably similar to that of Spirula
ballasting of the posterior end and a device for
(Figs 27, 28), though less strongly curved,shorter,with
burrowing (p 43).
5. The decreasingcurvatureis made possibleby the
a less inflated protoconchand more closely spaced
The peculiaritiesare closely connected with one
septa.The sheath posteriorly gradesinto a massive
another;it is unacceptableto declareone ofthem as the
rostrum, which surrounds the initial parts of the
cause of the others. Together they reflect a shift o./
phragmocone,forming a rounded elevation,named the
equilibrium in the whole organisation,which opened
"capitulum", around the protoconch.Anteriorly the
vp new habitats in relation to particular life styles and
periostracumbecomesprogressivelythinner; but on
thus generatednew pathwaysand perspectives(47) for
either side it appearsreinforced by the longitudinal
further modifications(see the closing sectionof this
"lateral swellings", which seem to represent the
part,p. 94). Here one point hasto be emphasized:
insertion sites of lhe musctrlarmantle.More ventrally,
During swimming, Orthoceras,the belemnitesand
the periostracumappearsparticularly delicate, a mere
the teuthoids have to achieve their position of
reinforcing layer of the conotheca;both together will
equilibrium by means of muscular effort. It seems
herebe calledthe "ventral wall" of the shell.
inconceivablethat a stable position (against rotation)
(46) The insertion of such a shell in the mantle sac
was achievedby the minimal excessweight provided
can be imaginedby the positioningof the phragmocone
by the ventral position of the siphuncle.Moreover, to
in a situation similar to that observed in a young
achieve a horizontal body orientation during
Spirula 6ig. 9). Posteriorly the insertion of the muscular mantle is situated on the capitulum, from
swimming, the buoyancyof the posteriorend had to be compensated,which resultedin partial cancellationof
where it continuesanteriorlyalong the lateral swellings
the original advantage and therefore in a waste of
to reach the free edges of the pro-ostracum,as is
energy.In the sepioid shell type, the gas content is
typical for decapods.This provides a prototype for the
shifted anteriorly and upwards, which makes the
derivation of all the sepioid shell and mantle shapes.
equilibrium state coincide with the normal orientation
How deep a hiatus exists betweenthis type and the
ofthe body.
generaldecapodtype has been emphasizedabove(Figs
An analysisof growth lines (Fig. 11b) showsthat
1I and 62). Here we shall formulate the essential
the curvatureof the phragmoconeinvolves difficulties
aspectsresponsiblefor this difference:
for the growth of the rostrum: in the course ol its
1. The muscularmantle has shifted its insertion
developmentthe tip of the rostrum must continuallybe
sites from the free edgeof the conothecato the outside
tumed ventrally. To reorientatethe tip accordingto the
of the phragmocone,in fact to the sheathsurrounding
k i n e t i c l o n g i t u d i n aal x i s . a n o p p o s i t em o v e m e n ti s
the phragmocone.
necessary;indeed,the growth axis of the rostrum must sheath undergoes special
slowly be curved upwards. This can be achieved by
difJerentiations for the fixation of the musculature,
2.
Thus, the
excentricaccretionas long as the rostrum is short and
similar to what happensin vertebratebones; these
simple in shape,as presumedfor Protosepioides (but
differentiations take the form of simple longitudinal
seeFigs 19, 23, 24). W e shall call this phenomenonthe "regulationofthe longitudinalaxis" (seealso Fig.71). - Sincethe earlieststageslack a rostrum (Fig. 10d), its
bulgesor ledges("lateralbulges"). 3. This necessitates a shift ofthe shell aperture(rim of the conotheca)to the inside of the visceralcomplex,
first rudiment no longer lies at the apex of the
30 phragmocone,which facilitates the solution of the problem. As to the anatomicalmodillcationscaused by the shift of the phragmocone(Fig. 9), see Naef (1913,p. 458).A deepincisionin the visceralcomplex, causedby the ventral shell rnargin, turns out to be inevitable;its immediateeffect is the decoupling(or severing)of the typical genital ligament, which attachesthe gonad to the shell in the area of the siphuncularentry (Figs 3a and 62). This vestigeof an (ontogenetically)prirnary mesentery disappearsin typical sepioids(Figs 9, 11) so that the gonadis only attachedto one side of the stomach. (48) The typical architectureof the other external and internal sofl parts and their primary correlations c a n b e t a k e n f r o r n F i g . 1l . ( B u t s e e a l s o t h e forthcoming critical presentationin Cephalopoda,vol. I, chapter38). It is noteworlhy that strong similarity to the teuthoidsexistsin many organs(Fig. 39). e.g.in the arm complex, which is surprising given the great difference in shell structure; this can only be
-l
I
i.tu.
understoodif we considerit an ancientheritage(seethe
1 f ,
closing section!).For example,the differentiationof
I
the tentaculararms is strikingly similar in all the living
Fig. 12. - Belentnosellaantericana (Meyer and Aldrich).
decapods(Cephalopoda, vol. I, p. I 15, 119),in contrast
Reconstructedfrom the original authors' figures (Pl. 2, Fig.
(Fig. 91). to thebelernnoids In the following sectionswe give a complete(as far as that is possible)descriptionof the sepioid.families,
26 and26a). (My reconstructionsshown by dotted lincs); nat. size. Compare Figs I I and 14 and note the undiff'erentiated stateof the capitulumand of the rostn.rm,the lengtheningand sharpeningof the latter, and the rounded lateral bulges. The
genera and species; it will be rather short due to the
small figures above the main figure shov' Belemnosis
scarcity of the material. We will take account of a
onomala(p. 52) for comparison(nat. size).
number of good descriptionsand illustrationsprovided by earlier authors, who may have missed certain offer invaluable details (v. Koenen, Cossmann,Meyer
Although the forms to be placed here doubtless representtrr,reSepioidea,in all aspectsthey - more
and Aldrich). Only the Sepiidae,which containmany
than other forms
speciesof uncertainstatus,will be treatedcursorilv,
Belemnoidea,as indicatedby the diagnosisand Figure
sincethey offer little in the way of new insightsand are
12. In terms of nomenclatureof the family, the genus
of limited significancefor the derivationof the group.
Belemnosisis typical; but in morphologic-systematic
generalrelationships,while their specialobservations
still resemble the older
terms the new genus Belemnosella shows more primitive conditions. D. The family Belemnosidae Naef l92l (System,p. 536)
1. The genusBelemnosellanov. gen.
Diagnosis: Sepioideawith only slight curvatureof the
Among the fossil sepioidsI have only recently found a
phragmoconeand inconspicuouscapitulurr, with short rostrum, - in which the insertionsof the muscular
form (Fig. 12) which in all its essentialparts (49)
mantle on the sheath [l.e. the rostrum] form rounded
group which I had already postulated.The earlier
lateral swellings,in other words they are not prominent
representation which
a n c u l a or r w i n s - l i k ee l e v a t i o n s .
morphological abstractionthus proved surprisingly
correspondswith the hypothetical prototype of the was
gained through
1 1
J,t
conect (cf. Cephalopoda, vol. I, chapter38). Not long
b.
ago considerationsrelated to Belemnosiscossmanni, describedbelow (p. 51), led me to slightly modify my earlierreconstruction,so as to be able to considerit the rr.rostprimitive sepioid (cf. Naef 1921, p. 540, and below Fig. l4). I was not then awareof the publication b y M e y e r a n d A l d r i c h ( 1 8 8 6 ) w h i c h c o n t a i n st h e illustration here restored as Fig. 12 and a brief characterizationwhich I have further interoreted rnorphologicallygiven the obviousrelationship.
f
Here I consideronly one species,known from only one specimen, identified by "BeIemnosis",namely:
the authors as
Fig. 13. Spirulirostrella szainochae(Vojcik) in dorsal (a), ventral (b) and lateral (c) views. nat. size. The rostrum has
Belemnosellaameficana (Meyer and Aldrich 1886). This form from the Tertiary (Eocene)of Missouri tWautubbee)differs markedly from the typical aspect
been added (dotted lines), in c the phragmoconeis also added. The protoconchprobably lies ventrally inside the thickened part anterior to the broken rostrum.
of the related genusBelemnosls,thereforeit cannotbe includedin the latter. In particularthe elongationolthe rostrum, and its pointed form must be emphasized,so
belongsto the Belernnosidae,being rather close to the
that this form seems to approach "Spimlirostra"
previousspecies.
s:ainochae (p. 50). A delicateslit startingfrom the posterior end of the ventral wall and cutting into the
Spirulirostrella szainochae(Vojcik 1903).
barely recognizable capitulum enhances this
Herebelongs"Spirulirostra" szainochae Vojcik 1903(1904), p.801-80P 3 ,l . 1 7F. i g .3 2 .A b e l( 1 9 1 6p, . 1 6 9 ) .
irnpression.(Could the shapeof the terminal tip, which has been reconstructedby the authorsof both species,
We are looking at a type that differs markedly from
be even more similar? Whatever the answer. there
Spirulirostra: althoughthe dorsalview showsa general
seemsto be no reasonto assumea marked difference).
aspectlike that of ,S.hoernesi(cf. Fig. 23), the rostrum
Unfortunately the description is rather cursory (p. -17):"Phragmocone rather long, (50) straightla,with
view the paft ('capitulum') surroundingthe protoconch
horizontal sutures.Rostrum obtusely conical below,
appearscompressedand split. Lower Oligocene of
cluadrangularlyflattened above". [Note: The holotype
Przemysl.(Prior to new investigations,nothing definite
is more distinct from the rest of the sheath.In ventral
of Belemnosella americona was redescribedand
can be statedabout this fossii. Probably we have a
refiguredby Jeletzky, 1969. According to Jeletzky the
Belemnosis-Iikeshell with a terminal spike (?), in any
type locality Wautubee is in Clarke County,
casesituatedoutsidethe seriesof Seoia\.
Mississippi,not in Missoury {sic} as statedby Naef. Jeletzky considered{1966, p. 106} that Naef's
3. The genusBelemnosisEdwards 1849.
reconstructionof a pro-ostracumin this specieswas
( c f . Z i t t e l1 8 8 5 p , . 5 0 9 ,F i s c h e r1 8 8 7 p, . 3 5 9 ) .
rvrong]. Belemnosiswhich is closeto Belemnosella,is the least 2. The genusSpirulirostrelluNaef l92l
specialized among the fossil Sepioidea. From
(System,p. 537 [536] and 541).
Belemnosella(5f it differs only by the obtuseshapeof the rostrum (Fig. 1a). Otherwiseit could pass as the
Here should be included the fossil describedbelow,
prototype of the whole group (cf. p. 49).However, the
which was originally identified as Spirulirostra.
original figures for B. anomala (p. 52) given by
Notwithstanding an overall similarity to the latter, its
Edwards (or Sowerby) do not necessarilysupport this
whole aspect is in fact so different that it must be
view. The peculiar aspect of this fossil from the
consideredas a representativeof a new genus that
London clay of Highgate (Eocene) is so striking,
)./. Fig. l,l.
Belentnosiscossmanni.Reconstructedfrom l/1 Cossmann. nat. size. - Original specimen in the "Bourdot collection".A fossil sepioidwhich
as far as it rs
preserved.and disrcgardingthe blunt rostrum
shows all
the f-eatures of thc ancestralform (Fig. 6f . a. Lateral view. The ventral wall (6) of the phragmocone. the outlinc of the mantle (11), and the parts of the shell above the brcak (,r) have been reconstructed. the phragmocone insidc the rostrllm has been inferred accordingto b and from observationsincorporatedin Fig. 2 t . p .6 2 . b. Ventral view" the anterior part reconstructed.ln the posteriorpart. the greaterportion of thc vcntral wall of the phragmoconc is missing; the septa are broken off. Their sutures(4) arc still recognizable,however. c. Idealizedcrossscctionclose to the shell aperture.The insertionof thc muscularmantle (9) on the su'ellingsof the periostracum(5) has been reconstructed.to illustrate the integrationof the shell in the mantlesac. 1. pro-ostracum;2. conotheca,dorsal part; 3. marginal thickcnings of the pro-ostracumat the transition from the conothecaas a continllationof J. 4. insertion of a septurn; 5. swellings of the pcriostracum for the insertion of the muscular mantle (9). 6. ventral wall of phragmocone (conothecawith thin periostracum);only some remarnsare preservcd; Z. "capitulum" of the sheathcontaining the "protoconch";8. blunt rostrum; 9. muscularrnantle; 10. siphuncle;11. outline of mantle; 12. integumentalcover. partsafter Cossmann(1895,Pl. l. Figs 3 and 4, Preserved p.164-165).
however,that it is reflectedin the speciesname. In
phragmocone almost to the summit, arched in the
contrast, the following, newly discovered form is
dorsal wall where the traces of septa are visible, with
typical in terms of its aspectand preservation.
two blunt edges on either side, and a hemispherical swelling in front of the last chamber" (p. 16a). "Blunt
a)
Belemnosiscossmanninov. spec.
summit, rounded, and freshly decayed" (p. 165).
(cf."B. anomala"Cossmann, 1895"p. 164-165, Pl. I. Figs3, 40). 4 , a n dN a e f1 9 2 1p. . 5 3 0 5
b) Belemnosis anomala (Sowerby 1829).
The fossil identified as "8. anomala" by Cossmann
Here belong, among others:BelernnitesanomalusSow. 1829
(52) certarnlyrepresentsa speciesof its own. The
(v. 6, p. 183,Pl. 59. Fig. 2); Belopteruonomalad'Orb. 1839
author gave a good illustration and description of the
( P l . 2 0 , F i g s 1 3 - 1 5 .1 8 4 5 ( p . 3 0 9 , P l . l z l , F i g s 8 - 1 0 ) a n d
specimen, but he seems to have missed its
onomala Quenstedtl8a9 (p. 4'73,PL.30,Fig. 1l). Belentno.si.s Gray 1849(p 158);Chenu 1859(p. 53, Fig. 167):B. plic'ata
morphological peculiarity. This peculiarity clearly showsup only when a reconstructionis attempted(Fig. 14), whereasa purely descriptiveconsiderationreveals
Edwards1849(p.,10,PI.2, Fig. 3); Edu''ards and Wood 1877, K e f - c r s t e i1n8 6 6( p . 1 4 3 9 ,P l . 1 3 0 ,F i g . l 3 ; B u l l c n 1 8 9 1( p .
no striking features.I quote essentialpoints from
289): Spirtrlirosrraplicata Bullen [Newton] and Harris 1894 ( v . I , p . 1 2 3 ) ; A b e l 1 9 1 6 ( p . 1 5 8 , 1 5 9 ) .( S c e a l s o B i i l o w .
Cossmann'sdescription:"Phragmoconefeebly coiled
1920,p.237-268).
in its posterior extremity". "Anterior part dilated".
According to Bullen fNewton] and Harris (1894)
"Horizontal septa"."Envelope decayed,with triangular
Belemnosis
incision in the ventral wal1, thus exposing the
Spirulirostra.
anomala
was merely
a corroded
This idea is correct as far as the
aa JJ
Fig. 15, - a) Belemnosisanomala from the London Clay of Highgate(Eocene),after Edwards(1849). a. ventral view; a1. )11 lateral view, nat. size. The most striking feature is the opening of the protoconch, probably due to post-mortem conosionofthe capitulum(cf. Fig. 12).
,W,
b) Plagioteuthis moscoyiensisRrimer and Dames (1890, p. 361, Figs 1, 3. 5) from the Oxfordian of Russia.An alleged"spirulid" with shor1,straight.blunt rostrum, which gradesdorsally into a
i"
VZ
strong ridge on the sheath.b. right lateral view, b1. dorsal vicw,
D2. anterior view of the aperture. The phragmocone (if it is one!) appearscompressed,u'hich would explain the asymmetric deibrmation of the "sheath" alluded to in the generic name. The "alveolus" is said to extend to the very end. The "sheath" is concentrically layeredicf. index,;.: r nat.size.
corrosion and the resulting opening up of the
64)
protoconch is concerned.A remote relationship with
1.The genusBelopteridium nov. gen.
Spirulirostra is undeniable.But identity with it (even at generic level) is out of the question,as emphasized
Here belongsFig. 16c; from the Paris calcaire grossier
a l r e a d y b y C o s s m a n n( 1 8 9 5 , p . 1 6 5 ) . T h i s a u t h o r
(Eocene)of Heronvalas:
indeed accepted the general (53) relationship:"The Belemnosis are Spiruliro.slrawithout a rostrum!and
Beloptefidium puerilis nov. spec.
with a less stronglycoiled phragmocone,which implies
I have seena hne, small specimenof this speciesin the
a clearly distinct genus;they neverthelessbelong to the
Bavarian State Collections (Munich
samefamily Belopteridae".
Collections); its quite indeterminatecharactermay
D'Orbigny thoughtin 1850(Prodr.2, p. 309) that it wasBelopter i na Ievesquci.
Public
explain its classificationwith the Beiemnitidae:the lateral edges(Fig. l6c) are totally blunt but continueas
A peculiar fact is the almost straight dorsal line
faint indications almost to the posterior end of the
when viewed in lateralaspect(a ); it curvessuddenlyat
rostrum. There they disappearin the region of the
the posterior end. This differs markedly from the
phragmocone;this region thus becomes strikingly
previous species;specificallyit indicatesa different
narrow. This difference from Belopterina is so
shapeof the posteriorparl of the body. Aparl frorn that,
conspicuousthat I have to createa new genus;its close
we see an overall aspect very similar to that of B.
relationshipwith the following one is obvious,
cossmannl.
however. One cannot exclude the possibility that this specimenis a juvenile form. It seemsconceivablethat the juvenile shell stagesof a Belopterina would look
E. The family Belopteridae s. restr.
rathersimilar.
N a e l 1 9 2 1( S y s t e mp, . 5 3 6 ) . 2. The genusBeloptefina Munier-Chalmas1812 Diagnosis: Sepioideawith a blunt rostrum that is lengthenedblock-like, the lateral insertions of the
This small group is in some respectsclose to
muscularmantle form blunt or sharplongitudinaledges
Belemnosis, (55,) whereas in others, especially with
or wing-like extensions(lateralwings).
regard to the shapeof the rostrum, it is closer to the
The peculiar differentiationof the massiverostrum,
gennsBeloptera, with which it was originally united.
which reachesits acme in the genusBeloptera (q. v.),
Both show a further but unequal strengthening and
suggeststhat the family represents"a deadlateralshoot
differentiation of the ridges which here replace the
on the sepioid stem". It indeed shows no close
lateral bulges of Belemnosl.r. From the Eocene
relationship to any extant type. Although the genus
(Paleocene). Nominal type:
Beloptera characterises the family, a few less specializedforms must be treatedhrst.
34 Fignre l6a and b representsa form of Belopterina which is clearly different from the previous one. I found the correspondingspecimen in the Bavarian State Collectionsat Munich (Public Collections).Of the alveolarpart only the posteriorpart is preserved, whereasthe ventral wall is lacking altogether.On the posteriorventral side also a piece is missing. The most characteristicfeatureis the form of the massive,nearly egg-shapedrostrum as well as the lateral edgeswhich appear to unite the Beloptera-like rostrum and the phragmocone;these edgesare clearly broader and are reminiscentof the jagged blade of a stoneknife. very Fig. f 6. - Different Belopterina-like rostra from the Eocene
different from .8. levesqttei. The gently incurving
of France, drawn at natural size. from specimer-rsin the
phragmoconeis easily recognizableand is relatedto
BavarianStateCollections(exhibitedcollection).
the curvature of the rostrum (see also Beloptera
a. Ventral view. b. Lateral view of B. deshoyesi(from Chenay near Reims).
edwardsiDeshayes1866,Pl. 107,Fig.3-4).
The ventral wa11of the alveolus is missing; it has been reconstructedin D. The shapeof the phragmoconeis shown
3. The genusBelopteraDeshayes1824?,Blainv. 1825.
by the depressionin the dorsalpart of the sheath.Thc rostmm is damagedventrally at the posterior end; the lateral ridges
T h e n a m eo f t h e g e n u s( B l a i n v i l l e1 8 2 5 ,p . 6 2 1 , P l . I I ,
are well preserved and much sharper than in the previous
in the original 12 by error) was given by Deshayesand
specles.
has been adoptedby Blainville, apparentlyby word of
c. -1.lateralview, 2. dorsalview of B. puerili,s(from Hdronval near Paris).Without distinctlateralridges. d. Lateralview ofB. levesquei(cf. Fig. l7). .rr.capitulum. containing the protoconch;_r. groove marking the courseof the mantle(fin) vesselsto the muscularmantle.
mouth. [Note: The genus was actually proposedby Deshayesin 18371.In Blainville's work it designates fossil Sepioideain general,namely our Belosepia and (56) Beloptera, the mutual relationship of which (as now demonstratedby intermediate forms) was thus r e c o g n i s e da l r e a d y b y B l a i n v i l l e a n d D e s h a y e s . Blainviile remarkson the similarity of Beloptera s. str.
a) Belopterina levesquei (d'Orb. 1839). The
original
diagnosis
of
the
species reads:
"appendicibus lateralibus parvis, linearibus"
Iwith
small, linear, lateral appendagesl (1. c.). The lateral wings of the better known Beloptera belemnitoidea (p. 56) are thus present as mere rudiments, which form angular ridge-like elevations. See Figures 16 and 11.
with belemnites (8. belemnitoidea) and of the other form with Sepio (8. sepioidea;cf . Belosepia,Fig. 33). Voltz (1836) also interpretedthem as intermediate forms betweenbelemnites and Sepia. We here restrict the scopeof the genus,thus following Voltz (1840) and (1872).It clearly joinsBelopterinarn Munier-Chalmas
Here bclongs:d'Orb. 1839,Belopteralevesquei.(Sepia
which the lateral edgeslink the rostrum and the conus,
( ! ) , P l . 2 0 , F i g s 1 0 - 1 2 ) ;l 8 4 l ( A n n a l c sp, . 1 9 ) ; 1 8 4 6
(Fig. 18) ( cf. Zittel 1885, formingwing-likeextensions
( P a l d o n t o l o g i eu n i v e r s e l l e . , P l . 8 . F i g s 8 - 1 0 ) ; 1 8 4 5
p. 509,Keferstein1866,p. 1439,Fischer1887,p. 259).
(Mollusquesvivants et fossiles,p. 308. Pl. 14, Figs 5-7);
Here belong severalspecies:
1 8 5 0( P r o d r o m e 2 , p . 3 0 9 , 3 3 8 ;Q u e n s t e d1t 8 4 9 ,P l . 3 0 . F i g . 40,Beloptera(.Beloprerina), Cossmann1913.Pl. 61, Fig. 3-2; Belopterina,Abel 1916,p. 142,Fig. 57. See alsoBeloptero Deshayes1866,Pl. 107,Fig. 1-2. leve.squei
b) Belopterina deshayesi E. Vincent 190 I . Here also belongs Beloptera (Belopterina) deshayesi Cossmann1913,Pl. 61, Figs 2-5, and Beloptera lcvcsqtrei DeshayesI 866,Pl. 106.Fig. 9-10.
a) Beloptera belemnitoidea Blv. Beloptera belemnitoidea Blainville1825.p. 622(Pl. II, Fig. 8 ; 1 8 2 7p. . 1 1 1( P l .I , F i g .3 ) ; d e S o w e r b1y 8 2 9( P l .5 9 ,F i g . 3); Sepiaparisiensisd'Orb. 1826,1839(Sepia.Pl. 3. Fig. 7p. l9); 9); Beloptera belemnitoidea d'Orb. 1841(Annales, ( P 1 . 3 0 , 1 8 5 0( P r o d r o m2e, p . 3 0 9 ) ;Q u e n s t e 1 8 4 9 F ig.88; dt Belopterabelemnitoidea Cossmann et Pissarol9l3 (Pl. 61, Fig.3-I);Belopterabelentnitoidea Leriche1906(Pl. 9, Fig.
35 6 ) ;A b e l1 9 1 6( F i g . 5 9p, . 1 4 4 ) . Blainville's original figure agreeswith those given in treatisesand textbooks(p. 58 below): b) Beloptera czrla Cossmann. Some of the specimensthat used to be placed in the abovespeciesli.e. Beloptera belemnitoidealhave to be distinguishedfrom it, a fact that led Cossmann(1896, P I . 2 ,F i g . 2 1 ,P I . 3 , F i g . 9 , 1 9 1 3P, l ; 6 1 ,F i g . 3 - 4 )t o create a new species, characterizedby: a) a short, poorly differentiatedrostrum, b) a blunt phragmocone, c) broad lateralwings, demarcatedfrom the rostrum by notches,which almostreachthe posteriorend. (57) Beloptera curta apparently was widely distributed in the Eocene (Paleocene)of France. According to Blainville it occurs in the Paris area; I have seen specimensfrom Beauves(Oise) and Ndhou (Manche). One known form has to be distinguishedfrom this species,and I thereforegive it a speciesname of its own:
Fig. 17. Belopterina levesquei(d'Orb.) reconstnrctedfrom 2/1 a specimenin the Bavarian State Collections at Munich. nat. s1ze. a. Lateral view of the shell fragment drawn in a three-
c) Beloptera longa nov. spec.
dimensional presentation,completed by the addition of
Only the original specimen of Figs lSa-g and 19
outline and soft parts.An essentialelementwas the formation
belongshere with certainty;possibly also some earlier
of the lateralridges.i.e. the insertionsof the muscularmantle,
examples,with indistinguishablefeatures,treatedunder a). See, for example,the median sectionin Edwards andWood 1877,PI.2, Fig. I. (58) My figures are basedon an exceptionallywell
in comparison wtth Beloptera (Fig. 19). The insertion of typical sepioid 1'rnswas inferred on the basis of the blood vessels leading to the fins. These can only have a course similar to that observedin Beloptera (Fig. 18c) where they have left more distinct impressionsthan in the present less
preservedspecimenfrom Bois Gouet near Nantes.lt is
differentiatedcase.The anteriorpart ofthe body in the figure
distinguishablefrom B. belemnitoideaby a) a longer,
shows typical sepioid features.Differenceswhich may have
more differentiated, boot-shapedrostrum, b) a more
existedcan no longerbe verifred.
slender, apparently longer phragmocone,c) narrower
D. Median section through the posterior part of the body.
wings which apparentlytaper towards the rostrum and
Note the cur-vatureof the phragmocone,the position of the
phragmoconesheath.The dorsal side is markedly roof-
rostrum in relation to it, and the bulge above the protoconch
like, with a median longitudinal ridge; the wings have sharpedges. I cannot refrain from considering the possibility that B. curta and B. longa might be the female and the
correspondingto the "capitulum" in Belentnosls(p. 5 1). c. Transverse section through the middle portion of the posteriorpart ofthe body, showingthe muscularinsertions. Note the hypotheticalcontinuationofthe lateral ridges as the margins ofthe pro-ostracumand compareFigs l4c and 21d.
male of the same species. As far as the overall
Pr. pro-ostracum; Sfr. lateral ridgc; Vl. ventral rlb; Wl.
proportions are concerned,this would correspondto
capitulum; Ro. rostrum; Efr. protoconch; Sw. septum; Si.
the situation observedin shells of recentSepia (q. v.),
siphuncle. Eocene (Paleocene)of France (Aisne-Lamotte,
in which the females show more or less markedly
Gilocoufi {Oise}).
broadenend lateral parts and a stouter bulge (phragmocone)than the males. In the present case, however,the differencesare not limited to a difference
Some of the specimensof Beloptera describedor
in the length-width relationship,so they should be
illustratedin the literaturecannotbe securelyidentified
features. consideredspecies-specific
to species.This is in particular the case for B. postera
36
'\q
,W
/w
L#t -\"----
\--\''2 --\-'->;
\2'--
,ffi)
F i g . 1 ; A b e l 1 9 1 6 ,p . 1 4 4 ,F i g . 5 8 ; v . B i i l o w 1 9 2 0 , p . 2 t 5 - ?1 7 \ ( 5 9 ) T h e d o r s a l v i e w o f a s p e c i m e no f " 8 . belemnitoidea" rn Edwards and Wood (1871, Pl. 2) showsconditionsdifferent from all the other examples. The specimenlooks much narrower than Fig. l8b and might representa different species,which could be named"Belopteraangusta"nov. sp.
4.
The genus Belopterella
Naef
1921
( S y s t e rpn. 5 3 6 ,5 4 1 ) .
\l/
'7111\ l---
"&N$
d)
Fig. 18. Shellsofthe genusBeloptera.drawn at naturalsizc from specimensin the Bavarian State Collections(public coliections)in Munich. a-g: B. longa nov. spec.;h-i: B. ctu'taCossn-r. and Piss. ru.ventral view; D. lateral view; c. dorsal view; d. transverse
Although clearly a member of the same family, the following speciesdiffers so markedly from the other speciesthat the authorconsidersit to lie outsidethe old genusBeloptera'.
Belopterellacylindrica (v. Koenen 1885). Here belongs"Beloptera" cylindrica v. Koenen, Pl. 4, p. 81, Fig. la-e. The illustrationis accompaniedby a good description,which is quotedbelow: "A single specimen is available; its preserved
section at the broken anterior end; e. cross section at the
length is 6 mm, to the beginning of the alveolus; it is
broadestposition;.f cross section of rostluml g. interior of phragmocone(dorsalvie*'); /l. ventral view; i. lateral view of
much more cylindrical (disregarding the wings and
h.
Edwards,Deshayes,Cornet and Briart from the English
their long insertions)than the speciesdescribedby and French Eoceneand from the Calcairede Mons. At the lower end the sheathis rather evenly rounded and
v . K o e n e n1 8 9 2( P 1 .1 0 1 ,F i g . 1 2 a - c p , . l4l4). This
has a diameterof about 1.5 mm, seenfrom below about
specimen is very incomplete; it certainly does not
2 mm, close to the alveolus about 2.3 mm (from front
justify creation of a separatespecies.I have been able
to back) and 3.8 mm when measuredacrossthe wings,
to examine the specimenin the Berlin Museum of
which extend only slightly more anteriorly (60) than
Natural History: conusand wings are missing; only the
posteriorly and which, at the level of the alveolus,are
rostrumwith the alveolarbaseis preserved.Theseparls
separatedfiom the feebly arched ventral side by
exhibit featuresof B. belemnitoidea(collection site:
shallow depressions,whereasthey are flat towards the
WesteregelnnearMagdeburg).
bluntly roundeddorsalside. On the mid-ventralsurface
The figures in handbooksand textbooks (Chenu
lies a delicateslirlike furrow which disappearstowards
1 8 5 9 ,p . 5 1 , F i g . 1 6 3 Z r t t e l 1 8 8 5 ,p . 5 0 9 , F i g . 7 0 0 ;
the lower end. The sculptureis finely warty, especially
K e f e r s t e i n1 8 6 6 ,P l . 1 3 0 ,F i g s 1 7 , 1 8 ;F i s c h e r1 8 8 7 ,P l .
distinctly towards the lower end". Occurrence:Eocene
2,Frg. 7) are generallybasedon Deshayes18241s(?)
nearCopenhagen.
(Pl. 100, Figs 4-6) and doubtlessrepresentBeloptera
A striking featureis the shapeof the rostrum which
(SeealsoGuettard1783,v. 5, Pl. 2, Fig. belemnitoides.
appearsrather undifferentiatedfor a belopterid;it is
1 1 - 1 2 ;d ' O r b i g n y1 8 3 9 ,S e p i a , P | . 2 4 , F i g . 1 2 , a n d
even lessdifferentiatedthan in Belopterinawhereasthe
1 8 4 5 ,P l . 1 4 , F i g s l - 3 ; C o s s m a n n1 8 9 2 ,C a t . E o c . ,p .
lateral edgesmust have formed more or less distinct
1 2 ; B u l l e n a n d H a r r i s 1 8 9 5 ,p . 1 2 2 ; C o s s m a n n1 8 5 5 ,
wings. In any case,this form clearly standsapar1.
Loire inf., p. 165,Pl. I, Fig. 1-2; Cossmannand Pissaro 1 9 0 0 ,p . 5 , P l . I , F i g . 4 , a n d 1 9 0 3 ,s u p p l .P . 6 7 , P L 1 4 ,
-
37 Fig. 19.
Beloptera longa, reconstructed from a
specimenin the Bavarian State Collections(Munich). naturalsize. a. Median section. b. Ventral view of the fragment,completed(dottedlines) and placedin its relationto the mantle. r'. Transversesection,showing the lateral wings at their greatestwidth and the insertionof the muscularmantle. r/. Transversesection of the shell in the middle of the phragmocone. c. Transverse section at the anterior end of the phragmocone,i.e. of the availablefragment. Note the relationshipof the phragmoconeto the rostrum, the structureof the phragmoconesheath,the position of the lateral wings in relation to both, the situation of the shell aperturein the soft body, the anteriorly decreasing curvature of the phragmocone,the distinctive shape of the rostrum. the position of the supposed"terminal organ", the courseof the mantlevesselsto the mantle 1cf. Fig. l8b) and the supposedposition of the stellate ganglia. Bl. buccal funnel; l/. outer 1ip; 1/. inner lip; Ofr. upper mandible; Zg. tongue; Ob. upper buccal ganglion; U&. lower buccal ganglion; Cg. cerebralganglion;Nft. collar attachment; Sd. poison ("salivary") gland; Pr. proostraum (may be narrower!); Pi. phragmocone;Or. terminal organ (hypothettcal); Mm. muscular mantle; Hz. hear-t;Nl. kidney; ID. ink sac;Ed. intestine;Ro. rostrum; Co1,Cd2,Co3.coelomic sections;Ov. ovary;B/. caecum; -l1a' stomach;Gg. stomachganglion;Ao. aorta;Lb. Iiver; Oe. oesophagus;Zc. vena cava; Vg. visceral ganglion; Sl. statocyst;pg. pedal ganglion;Dr. funnel gland; Ik. funnel valve; Sr. subradularganglion; Ufr. lower mandible;Sl. siphuncle;,Sr,r.,. septa;S/r.lateral ridges; Sp. lateral wings; Eft. protoconch.;Mh. mantle cavity; Mrn. muscular mantle; Sr. stellate ganglion; pr. pro_ostracum;Kb. branchialband (insertion on the mantle); Vl.Yena pallialis lateralis (entry to the mantle); Vp.Yenapallialis posterior (ibid.); lp. afteriapallialisposterior.cf. Cephalopoda,vol. I, TextFig.265 and explanations.
F. The family BelosepiellidaeNaef 192I
lateral wings at the posterior end of a Beloptera shell
(System,p. 536).
once the club-shaped rostrum has completely (cf. Abel 1916,p. 146,Fig. 60). According degenerated
Only one genusis consideredhere:
to Leriche (1906), the Dentalium-ltke shell of Vasseuria Mun.-Chalmas would represent the
The genusBelosepielluAlessandri1905.
phragmocone belonging to
Two forms from the Middle Eocenebelong here; their
agreement.The shellsof Vasseuriaare the sheaths[i.e.
sepioid characteris somewhatdoubtful. I nevertheless
rostra] of belemnite-like shells, as will be shown
considerthis interpretationappropriatefor the present.
below; they are by no meansphragmoconesthat could
The possibly identical speciesB. cossmanni and B.
be combined with the shell of Belosepiella(cf. Fig.
parisiensisAless.(1905,p. 146)are limpet-likeshells
e6).
(Fig. 96) showing a gap on the convex side as if a Beloptera-like phragmoconehad been inserted there. One can visualize its formation by uniting a pair of
B . This seems i n c o n c e i v a b l e ,i n s p i t e o f A b e l ' s ( 1 9 1 6 , p . 1 4 5 )
38 phragrnocone ventrally incurved and very large capitulurn,- in which the lateral edgesare prominent as lateral flanges,originating from the capitulum, that extend to the base of the pro-ostracum,on the lateral marginsof which they taperoff.
t
/r
c a.
@ b.
c.
d.
1. The genusSpirulirostridiumnov. gen. We begin to consider the series of spirulirostrids, which appearsto approachthe sepiidtype in a stepwise fashion,by looking at a form that is closely related to Belemnosis'.
e.
Fig. 20. - Belopterella cylindrica (v. Koenen).Drawn fiom 2/r v. Koenen(18S5,Pl. 4, Fig. 1a-c)at naturalsize(a) or at
This speciesoccurs in the cementstonesof the
nat. size (D-e).Theonly preservedrostrum with the posterior
Lower Oligoceneof HAring near Kufstein (Tyrol). I
end of the alveolus is drawn in three dimensions, the
found the original specirren (labeled "Beloptera
remainingparts are rcconstructed.The belopteridcharacteris c1ear,but the rostrum is less specializedthan in the other forms. The lateral wings are intermediate in their developmentbet\\''eenB eloptera andB elopterin a.
Spirulirostridium oblusam nov. spec.
spec.") in the Bavarian State Collections (exhibited collection) in Munich (Fig. 21). Professor Dr. Schlosserkindly provided two additional specrmens
fracturc surface, showing the posterior part of the alveolus
from the sarnelocality for examination.They show the sameconditions,as far as my observationsgo. - In the
and the latcralridges.
unpreparedstate the original specimenonly permitted
D. dorsal, c. ventral. d. lateral'view, e. anterior view of
recognition of the overall shapeof the posterior end, which strikingly resemblesBelemnosls, on the one G. The family SpirulirostridaeNaef 1921
hand, and Spirulirostra, on the other, thus making it
(Systern, p. 536).
most interesting to me. With the kind permission of Professor Broili I have prepared the shell and (after
Much more interesting conditions are observed in a
drawing it carefully) ground it to the median plane.
seriesof fossil speciesthat can be united in the farnily
Thus,the structuresshownin Fig.21 were exposed.
Spirulirostridae.Although they are known in Europe
This shell quite ideally fits in between Belemnosis
only from the Middle Oligoceneand Miocene, they
cossmanni (p. 51) and Spirulirostrahellardii (p. 63),
show the general aspect that one would expect of
thereforeat first I consideredit an essentiallink in the
preliminary stagesof the actual Sepia type, which
Seplaseries.
occurs there already in the Eocene (61) (Belosepia).
(62) However, study of Belemnosellaamericana (p.
However, the Eocene sepiids had very strong, larger
49) refuted this view; I now tend to considerboth
shells (about 20 cm); but except in a few rare cases
Belemnosisand Spirtrlirostriditrm as secondaryforms,
only the strongestparts of them are preserved;in
the former derived from Belemnosella,the latter from
contrast,the spirulirostridshad small, delicateshells,
Spirulirostra. If the earliestparts of the sheath(Fig.
a n d t h e i r f o s s i l i l i z a t i o n p r e s u p p o s e sp a r t i c u l a r l y
22b), as I suppose,show an indifferent form without a
favourableconditions.N evertheless,spirulirostrids are
sharp rostrum, the formation of a rostrum may have
known from the Eoceneof Australia (Fig. 23d-f); they
been inhibited altogether, as apparently occurred in
show us how prematureit would be to concludefrom
Spirula (Fig. 26) (cf. Fig. 21). Ar presentI think that
the present absenceof a type in our material that a
the pointed rostrum of Belemno^sel/arepresentsa
morphologically substantiated relationship is
heritage frorn the (older) belemnoids and can be
impossible.(The same is the casewtth Spirulirostrina
supposedto have been conservedin the lineage of
at the presenttime).
S p i r t r l i r o s t r a , r v h i c h d o u b t l e s sp r o v i d e s ( t h i s i s
Diagnosis: Sepioideawith the posteriorpart of the
essential!)a more straightforwardunderstandingof the
39 Fig. 21. - Spirulirostridium obtusumnov. spec.(2/1 nat. size). Fossil sheil, from the Cement marl of Hiiring near Kufstein (Tyrol). Lower Oligocene. Prepared and partly ground down. Original specimen in the PalaeontologyMuseum (public collection)at Munich. a. Ventral view. The dottedparts of the outline, and the free margin of the phragmocone and its continuationas the pro-ostracum,are reconstructed. CompareFig. 14,p. 23.
n*?.8
b. Reconstructed median sectionof the posteriorpart
'r&
ofthe body.
I ii.
c. Specimenground down to a median section,the growth axis of the rostrum marked by a dotted line 1r)..t. remainingpart of the matrix. r/. The anteriorend of c ground to show a transverse section.The reconstmctedparts of the shell (which is only partly freed from matrix) are dotted. Integumentalcover (13) and muscularmantle (B) are indicated.
r5
c,. Lateral view. Same orientation as in c, anterior parts (dotted) reconstructed.The ventral wall of the phragmoconeat 20 addedfrom the sectionat c. (See also Cephalopoda, vol. I, Fig. 266).
Fig. 22.
a and b. Spirulirostra bellardii, posterior end reconstr'r.rcted from
the specimenshown in Fig. 24A: a in lateral view, b in median section.One can easily imagine these parts incorporatedin to Fig. 11. Indeed, the rcconstructionof the soft body must be based apart from the immediate effects of the shell on the structures- on the same principles, that is the recognition of the typical conditions in living sepioids. We cannot know anythingaboutpossibleatypicalpeculiaritiesofthe species.CompareFig. I I and its explanation,and there again note the upward deviation ofthe growth axis of the rostrum, the progressive tapering off, and the rugose structuresat the position of the capitulum,where the muscularmantle is inserted(cf. Fig. 24a).2l1nat. size. r'. Anterior view of a shell of Spirulirostra hoernesi v. K., drawn from a specimenin the Berlin Museum (not the original of Fig. 23a-c).The alveolus and the anteriorpart of the sheathare representedas cut across.(cf. Fig. 24 Ba and note the striking width of the shell).Nat. size.Locality: Dingden near Mrinster.Miocene.
40 The genus Spirulirostra is closer to the recent sepioids than any of the previous genera; for morphological reasons
explained
in
my
Naples
monograph
(Cephalopoda, vol. I, chapter 38), (63) it might be the prototype of all the living sepioids. Of special interest is the fact that the genus contains several species which,
from
a state reminiscent of Belemnosella,
appear to gradually approach the derived sepiid type.
a) Spirulirosta
bellardii d'Orb. 1842.
(cf. Bellardi,Moll. Ter. Terz. Del Piemontee Liguria I, p. 1 9 .P l . 2 , F i g .8 ) Sp.bellardiid'Orb. 1842(Ann.,p. 362,P1.11,Figs l-6). Sp.bellardiid'Orb. 1842(Pal.univ.,Pl. 9. p. 295). S p . b e l l a r d i i d ' O r b . 1 8 4 5 ( M o l l . v i v . f o s s . ,p . 3 1 1 , P l . 1 5 , Figs I -4).
l$ i r:{3{ ",J'H
W,
Fig. 23.
a-c. Spirulirostra hoerne.siv. Koenen 1867 from the Miocene of Dingden and Berser-rbnick in Westphalia.3 1 nat. size. Reconstructedfrom thc figures and indicationsofv. Koenen. The broken apex of the rostrum is reconstructed,as
Sp.bellardii d'Orb. I 852 (Prodr.,vol. 3, p. 25). Sp.bellardii Quenst.1849(ibid. Gray, 1849). Sp.bellardiiChenu1859(p.52, Fig. 164,165). Sp. bellardii Kcferstein1866(p. 1439,Pl. 130,Fig. 19). Sp.bellardii Fischer1887(p. 358, Fig. 136 {after Mun.-Ch.. ParisMuseum|). Sp.bellardiiZiucl 1883(p. 510.Fig. 701). Sp.bellardii Pompeckij 1912(p. 296d, Fig. 57 {from Stromer
1e121). Sp.bellardiiAlessandri 1897(p.a8). Sp.bellardiiParona 1898(p.167,Pl.XII, Figs8-l l). Sp.bellardiiAbel1916(p 157,Fig.63etc).
is the pro-ostracum (dotted lines mark the fracture). The phragmocone is restored to its natural position in a, and
This speciesoccurs in the hills (Superga)near Turin
representedaccordinglyin D and c'.The reconstructionis thus
(Miocene);the original figures given by d'Orbigny are
of small extent and is based in every respect on secure
essentialfor its characterization.They illustrate a form
observations.As to the position of the shell in the animal, see
(61) from the samelocality that I have seenmyself (cf.
my Naples monographand Figs 22a and 11. Comparealso the original figuresofv. Koenenin Pl. 14, Fig. 6. 1. pro-ostracum:2. frec ventral apertureof conotheca;3. phragmocone (vcntral wall); 4. lateral wing; 5. capitulum, containingthe protoconch;6. rostrum; 7. last septum;B. free margin of conotheca,dorsalpafi seenfrom inside.- a. lateral view, D.dorsalview, c. ventralview. d-f. Lateral, dorsal and ventral views of Spirulirostra curta from the AustralianEoceneafter Tate (1893, Pl. l, Fig. l). r/4 About nat. size.
Fig. 24A). In this form the protoconchis embeddedin a thick, rounded knob of periostracumthat offers only some inconspicuousrugose structuresfor the insertion of the muscularmantle. b) Spirulirostra hoernesi v. Koenen. c f . v . K o e n e n1 8 6 5p, . 4 2 8 1 8 6 7p. . 1 4 5 - 1 4 6 P,l . 1 4 ,F i g .6 : 188);p.352-3s3. Koenen justly considers this species, although doubtlessclose to Sp. bellardii, as a separatespecies. The detaileddescriptionindeedarguesin favour ofthis
whole diversity.Spirulirostridittm would thus represent
position, and after personalexaminationI came to the
a diverging branch of the line which leads, via
conclusionthat it certainly is a well defined, separate
Spirulirostra,to the sepiids.
type (Fig. 23a-c).I quote from the original description (v. Koenen 1867,p. 146): "The chamberedalveolusr6
2. The genusSpirulirostra d'Orb. 1842.
lbnns an almost completewhorl; (65l I was not able to
41
6 L/
..7
Comparisonof Spirulirostra bellardii (A) and Sp.sepioidea(B), drawn from fragmentsfrom Turin, now in the Bavarian 2 StateCollections(publiccollection)in Munich. I nat.slze. A1. B. Lateral views liom the left side; reconstructedparts dotted. ,A2,82. Median sections;82 drawn from the split specimenA.2, Fig,24.
completed by adding the integumentalcovering, insertion of muscular mantle, and coelomic epithelium. A:, B:. Ventral views, reconstluctedpartsdotted.Ba. Anterior view of B, to show the capitulum(cp), the ventralprocessbelow it, the lateralridges (.ry),the ventralwall with rib (Vr),the position of the siphuncle(si) and the tbm of the transversesection(cf. Fig. 22c). cp. capitulum;sp. lateral ridge; vl ventral process;pr. pro-ostracum,in reality much longer (cf. Fig. 23a); x. apical line, i.e. growth axis of rosflum;-r. growth line on mediansection. A comparisonbetweenthesepicturesis interesting:the capitulum in A is more massive,simply rounded,broad,surfaceonly slightly rugose(cf. Fig. 22b) in the position ofthe ventralprocess.
examine its end". As to the median sectionthe author
"The straight portion of the alveolus is preserved
remarks:"in the earliestjuvenile stagethe outer shelllT
only in the specimen figured; it rapidly increasesin
merely forms a blunt protuberanceimmediatelybelow
thickness and finally measures10 mml8, after having
the spiral alveolus but then rapidly (6 mm without
attained 5 mm in diameter; its length is 18 mm. The
check) grows out to a point, subsequentlyadding thrn,
septaare about 2 mm apaft in the terminal parl, about I
concentric envelopes.From the tip to the alveolus
mm in the earlier sections.The whorl of the alveolus
extendsa distinct "midline" similar to that observedin
measuresabout 7 mm, its beginning about I mm in
belemnites, and on a transversefracture a radial
diameter.The cross sectionof the alveolusis perfectly
structureis also recognizable.The surfaceof the outer
circular throughout". The specimenswere collected
shell is covered with warty granules, sirrilar to the
from the Miocene of Westphalia near Dinsden and
surface of the cuttlebone; these granules are sharply
Berssenbriick.
delimited on their inner side, and the inner shell layers
A difference from Spinrlirostra bellardii, according
likewise are made up of distinct granules, so that in
to the author(1882,p.352), is "a stouterform, much
sectionsa granularconglomerateappears.
shorter tip below and more strongly curved alveolus.
42 The latter makes up about one third to three quarlers of
my new species rather than Spirulirostra bellardii
a whorl, apparentlybeginningwith a globular chamber
d'Orb.; thus its occurrence at Turin is further
about 0.5 mm in diameter.Unfortunatelythe septaare
processusventralis is also confirmed.The nose-shaped
broken. The earliest septa lie about 0.75 mm apart,
visible in one of the specimensdescribedby v. Koenen
l a t e r t h e d i s t a n c eb e t w e e ns e p t ai n c r e a s e st o I m r n .
(P1.XIV, Fig. 6a); I thereforetransferthis forrn to my
finally to about 2 mm. The diameter of the alveolus
new species. From the co-occurrence one might
frrst measures0.5 mm and then increasesto nearly l0
concludethat,!p. sepioideamerely representsan earlier
mmrE". According to my observationsthe insertion of
stage of Sp. bellardii. This is contradictedby the fact
the lateral plates on the capitulum presentsa slightly different aspect(cf. Koenen 1865,Fig. 6h and my Fig.
that dp. sepioidea is f?barely] larger and more robust than ,!p. bellardii, as demonstrated(as far as is possible
22c with Fig. 24Ba). The lateral plates appearbroader
to date) by the specimensfrom Turin and Westphalia,
and are also wider in the posterior part. The juvenile
respectively.Doubtlessthe young animalswere lacking
shell thus appearsabout as broad as high (dorso-
the special featuresof the speciesso that they looked
ventrally) and the animal must have displayed a
more similar to Sp. bellardii of the same age than did
circular cross section in the mantle sac, insteadof the
the adults. But this is a generalrule valid throughout
laterally compressedoutline of Spirulirostra bellardii
organic nature.
and fp. sepioidea.The capitulum, in contrast,appears very naffow comparedto the (66) massive lorm seen in Sp. bellardil. - The other form found in the State
p. 167,Pl. 10,Fig.1;Chapmann Tate1893, 1914.Fig.106F.
Collectionsat Munich, which is describedbelow (Fig.
This species,which is apparently close to ,!p.
248), apparently differs little from the two above-
sepioidea (cf. Figs 23, 24), occurs in the Eoceneof
mentionedforms. However, it is noteworthy that it
Australia.We quotefrom the author'sdescription:
clearly approachesthe more modern sepioid types (Spirulirostrina, Belosepia,
Sepia)
by a mere
d) Spirulirostra curtu Tate 1893.
"Nicholson placedthe genusin Sepiadae,Fischerin Belopteridae,Tryon in Belemnitidae;this last location
modification of the samebasic form: a new elementof
seems to me to be the best, as Spirulirostra may be
the sheath appears completing the "dorsal shield"
viewed as a belemnite (68)
which replacesthe above-mentionedrugose structures
phragmocone lying obliquely within the alveolar
(Fig. 22b, cf. Fig. 24&). (67) This, as well as the
cavity". "The rostrum of the Australian speciesls more
modification of the overall proportions,is a reasonto
robust, is shorlly pointed, and less archeddorsally and
distinguisha separatespecles:
ventrally, where it is more or less truncated. Lateral
with a subspiral
axis of rostrum 8, ventro-dorsalaxis 12, length to apex c) Spirulirostra sepioideunov. spec.
of phragmocone16; length of alveolar cavity above
The type specimen is the fossil in the Munich
plane of phragmocone 29 (incomplete)". The
collectionsillustratedin Fig. 24B; accordingto the
measurementsgiven do not agree with the figure,
records it also comes from Turin. I call the newly
which is approximatelynatural size (my Fig. 23d-{). If
appearing structure, which has its hornologous
the frgure is correctly drawn, Sp. curta differs from ,!p.
structures,the processus ventralis. It seems initially
s e p i o i d e a b y a n i n c o m p l e t e d e v e l o p m e n to f t h e
quite independentof the lateralridges;however,on the
processus ventralis, by a stouter rostrum with less
sheathone flnds rugose structuresthat look as if they_
pronouncedlateral compression,which also affectsthe
indicate a true link with the ridges. Thus a secondary
correspondingmantle sac. Since the distancein place
continuous shell margin would also be formed at the
and time is also considerable,I consider Sp. curta a
posterior end for the insertion of the muscularmantle,
separate s p e c i e sT. h i s s e p a r a t i o ins i m p o r t a n ts i n c ei t
which would be perfectly comparableto that observed
confirms the Eocene age of the genus, which already
tn Sepia;at any rate it is necessaryfor the derivationof
had to be assumedtheoretically(p. 61 and 76). (For
the cuttleboneof Sepia(cf. Fig. 29).
Spirtrlirostra from the Oligocenealso see Sacco 1904,
Incidentally,this form is not entirely unknown: the specimendescribedand illustratedby Michelotti (Pl. 15, Fig. 2; cf. ZrtteI 1883,p. 50, Fig. 701a)represents
p. 5). The descriptionand illustrationsof Tate have becomeavailableto me only recently.
43 The genusHeliceras (Dana 1848)em. Fischer1887.
1
1
Heliceras /ilgensis Dana 1848 is an incompletely c h a r a c t e r i z e df o r m o f s e p i o i d , w h i c h m a y b e
h
mentionedhere as an appendix.Chenu (1859, p. 53) quotes the following diagnosis from Dana (whose original publication was not availableto me): "Animal unknown. Shell thick, subcylindrical,presentingon its inner side a compacttubular cavity at the extremity of an alveolus and ending in a chamberdivided by spiral septa". Fischer(1887,p. 465) usedthe samesource:"Shell similar to a belemnite;rostrum subcylindrical,thick, of fibrous texture; phragmoconedelicate, ending in a spindle-shaped,spiral nucleus"; "Collected in a slaty rock of CapeHorn."
H. The family Spirulidae(d'Orb. 1826)Owen 1836. Diagnosis: Sepioideawith phragmocones that have a circular cross section and oblique septa, that are spirally coiled and have only sparseremaindersof a sheathin the form of a crust-like cover,- in which the pro-ostracum has entirely (69) dtsappeared and is topographicallyreplaced by the muscular mantle, the dorsalshellmargin being pushedinsidethe mantlesac. D'Orbigny alreadyconsideredthe living Spirula as
Fig. 25. - Spirula spirula (L.). Adult female at natural size.
a representativeof a separatefamily ("Spiru1ees"),for
Drawn from a specimen in the Vienna Museum. slightly
which Owen later used the presentname. It is known
simplified, the missing right tentacular arm reconstructed.
sincethe Miocene from the fossil record;today it has a
Comparethe mantle inserlion (ir) on the shell in the juveniie
worldwide distribution in marine coastal sediments. After the deathof the animal the gas-filled shell (much like a sepiid cuttlebone)rises to the sea surfaceand is carriedaway by surfacecuffents.The fossil shellsvery
specimen of Fig. 26a. Although without any essential modification, the mantle sac appears to extend more markedly in a posterior direction on either side (,t) and to be insertedbetween the two indentationsof the terminal organ (l). The latter bears the fins, the slanting insertion lines of
much resemble the recent ones, and the same is
which are oriented dorso-ventrallv.entirelv indeoendentof
probablytrue with regardto the animal.
the muscularmantle. The specimenwas collected off German East Africa by an
The genusSpirula Lam. 1801.
officer (on board S.M.S. "Frundsberg"). who brought it to Vienna. It is said to have drifted at the surface.The skin is
Diagnosis: Spirulids with shells growing spirally, at least during the juvenile phase,finally reaching2t12
exposedin the "window" area; parts of the apparentlytough shell sac are preserved.d. position of the dorsal shell
whorls, the last one half whorl deviatingfrom the spiral
aperture;s/. supporting ridge of the penultimate septum; 1.
thus shou,inga tendencyto becomestraight(Fig. 23),
eye lens, largely covered by the primary lid, so that only a
in which the posterior end during the post-embryonic
narrow openingremains.
phasedevelopsa conical knob which is surroundedby a fleshy ring-shaped elevation, so that the whole
-=
eroded, as is usually the case, but the shell is not entirely
44 Fig. 26. - Half-grown Spirula at natural size (cf. C. Chun, 1915, p. 126, Fig. 36). Drawn from the original specimen and reconstructed using all available material, especially the specimensstudiedby Chun, his drawings and the earlier figures ofOwen (1878),Huxley (1895),and Lonnberg(1896).(Notethe Sepio Ia -llke habitus!).
nd
a. vcntral view. &. dorsal view. c. lateral view. - Fig. c'is Itok-
representedas transparent,so that the muscular mantle, ventral integument,anal papilla (an) andterminal organ are visible in an optical section, whereas the shell and some organs are
*
represented in lateralview (cf. Chun 19 I 0, Pl. I , Fig. I ). tS tentacularbaseinsidea simplepocket; ya. ventral insertionof the mantle on the shell; an oval areaof the latter is seenthrough
the thin integument.whereasthe posteriorpart (.r) is hidden by the terminal organ (lo). da. dorsalinsertionof the muscularmantle;1 asr! i'. iris fold; /l. cye lens. ld. primary lid. slightly contracted,whereas figures a and 6 show it open; ntd. dorsal,median section through the muscular mantlei lc. retractorcapitis; r.r,t.living chamber; da. dorsal insertion of the muscular mantle on the shell; ht. skin;fl. fin; st. terminal spine, retractedinside the tcrminal organ; r.l. ventral insertion of the muscular mantle; /r7s.mantle septum; ny. ventral median section throughthe muscularmantle; mh. mantle cavity' an. anal papilla; ki. glll: rt. funnel retractor;tft. funnel attachment;ll. funnel pouch; nh. collar attachment;r'o. olfactory organ.
structure("terminal organ") looks almost like a sucker,
therefore draw the reader's attention to the fine
which have (70) smal| roundedfins, similar to those
specimenof Fig. 25 which I was able to study at the
of sepiolids, which are obliquely inserted on the
Vienna Museum; after taking all the measurements,
posteriorend, the surfacebeing directedanteriorly and downwards,- in which the simple elongate funnel
somepartshavebeenreconstructed in l1rydrawing.
cartilage occupies the whole length of the funnel
d e v e l o p m e n ti s g i v e n i n t h e N a p l e s m o n o g r a p h
pouch,
A more detaileddescriptionof the animal and of its
in which the primary eyelid can be opened
(Cephalopoda, vol. I, chapter39). Here I merelygive a
widely throughout life (it is only temporarily closed,
summary and some figures, to make this living fossil
except for a small pore, when the eye needs to be protected),- the arms of which increasein length
among the dibranchiatesknown to the reader and to facilitate a better understandingof its fossil relatives.
slightly from the dorsal side downwards(arm formula:
Among other specimens,I had at my disposal the
4 , 3 , 2 , l ) , e a c ha r m h a v i n g4 r o w s o f s m a l l s u c k e r s
material, including juvenile stages,describedby Chun
with smooth rings, whereasthe tentacular clubs have
(1910,1915).
in which all the arm
(71) The shell of adult Spirula is well known and
bases are connected by a distinct interbrachial
well representedin museum collections (Fig. 26c).
membrane.
Appelldf (1893) describedthe fine structure,which is
their suckersin about 16 rows,
Fossil ,lplrzrlawere not availableto me. They occur
generally confirmed by our observations;our view
in the Miocene of Italyre.The recent forms all belong
differs only in terms of their morphological
t o o n es p e c i e sn.a m e l y :
interpretation. It should be emphasizedthat the shell of Spirula
Spirula spirula (L.1161) Hoyle 1909.
totally lacks a pro-ostracum,as in the shell of Sepia (q.
This speciesappearsto occur in all open seas(Atlantic,
v., and p. 4l). Otherwise all the typical parts of a
Pacific, Indian Ocean)and to occasionallyrise to the
decapod shell are present,in particular the ostracum,
sea surface.The normal habitat must be the cold deep
hypostracum, periostracum, septa, siphuncle,
waters where they apparentlyoccur in great numbers,
prosiphon. In the correlation of these composentsand
to judge from the massesof strandedshells. Well
in their particular form, we find a characteristic
preservedanimals are rare in museum collectionsz0;I
modulation, however; but behind these modifications
45 ,rne can easily recognize(Fig. 2b) the generalshell lr pe and the speciai relationship of form with .\pirulirostra. In the context of internal shells, a surprising .'haracteris the almost completespiral coiling. This iannot be explainedsimply by recalling the derivation trf nautilids from straight tetrabranchiates. There is an rrndeniablesimilarity betweenSpirula and Gyroceras. However,in the latter the early chambersturn upwards, rvhereasin Spirula they turn downwards. What is nrore,the coiling of Gvrocerasinvolves the whole soft body, which not only occupiesthe shell aperture,(72) as do all sepioids, but also accommodatesall the .juvenileparts. The loss of the pro-ostracumis probably relatedto the spiral coiling; indeeda pro-ostracumwould not be able to keep pace with the continuous spiral growth, nor could it be formed in direct continuationof the spiral. Likewise the disappearance of the rostrum is an inevitableside effect of this shell form. The secondary u n c o i l i n gt n B e l o p t e r a a n d S p i r u l i r o s l r a c o u l d b e relatedto an inhibiting effect of the rostrum; likewise the presenceof a rostrum would have inhibited the coiling of the shell in Spirula. Therefore the rostrum rnusthave been lost prior to the attainmentof the form
Fig, 27.
a. Median section through the initial part of a
Spirula shell. r2lr nat. size. This figure integratesthe data and figuresof Appellttf(1893),especiallyhis Pl. 9, Fig. 1.
The
primary shell wall (ostracumand hypostracum)is marked by cross hatching (6), the periostracumis dotted (7); as are the
of Spirula. The samecan be said of the lateralplates.
septaand the calcifiedseptalnecks(18, 15, 13,5, 4). In the
Although the sites for insertionof the muscularmantle
siphunclethe epithelium of the soft body is marked by dots.
are basically the same as in other sepioids,namely the
-1.protoconch; 2. prosiphon (strut); 2a. sagittal lamella of
sidesof the phragmocone,permanentlateral insertions
prosiphon; J. initial parl of the conchiolin cap which forms
are incompatible with a continually coiled shell
the first part of the siphuncle;4. adjoining septalneck of the
growth. Therefore the periostracummust be reduced,
(rudimentary)first septum; 5. conchiolin neck of the second
becoming limited to a crust-like envelope of the phragmocone;in other words, it returns to its original condition(cf. p. 175:Belemnoidea).
septum (extendingthrough l2 Io 13);6. shell wal1, ostracum and hypostracum; /. periostracumlB. initial part of fleshy siphuncle; 9. first septum and septal neck. ventral part; 10. enlargementof fleshy siphuncle; 11. pillars; 12. connecting
Spirula thus appearsboth conservativeand derived,
r i n g ( b e g i n n i n g ) ;1 J . s e c o n d s e p t a l n e c k ; 1 4 . s e c o n d
which gives it a clearly isolatedposition within the
chamber; 15. second septum; 16. angle deposit; 17. ptllar
suborder.One wonders how this modification of the
belonging to the third septalneck; 18. third septalneck; 19.
typical form may have been ecologicallyenhanced,i.e.
siphuncular epithelium; 20. thtrd gas chamber; 2J. ventral
what
advantages and
pan of third gas chamber;24. venrralrib of periostracum;25.
disadvantages, what effects the adaptivemodifications
second septal enlargementof siphuncle; 26. ventral part of
could have been i{s
might have had. Unfortunately a direct observationof the living animal was not possible,so we can only deduce the following points from
structural
considerations:l) The reduction of the pro-ostracum
secondair chamber. b. Hypothetical median section through a newly hatched juvenile animal.6/1nat. size.c1'.Figs 9 and 10. 1. muscular mantle; 2. remains of pro-ostracum;3. shell sac (she1l epithelium).
correspondsto the expansionof the muscular mantle (cf. above p. 24) and thus reflects a general developmentaltrend of dibranchiates.2) The coiling of
that correspondsto an essentiallyhovering, plankton-
the phragmoconeleadsto a reductionof the mantle sac
hunting animal which dependsmore on short dashing
46
---,0.
p0, 05+hy
b
this facilitatesa secondaryextensionof the mantle sac, P0 05
possibly (as in many other living forms) related to
v,
gonadmaturation. The shell wall (Fig. 28b) clearly shows,at leastin the later pafts, the typical three layers:The ostracumis thin, strongly refractive, the hypostracum is thicker, foliaceous like mother of pearl but without its typical gloss and not very refractive. The periostracum is a
0s+hy
thin bark-like, coarse,knobbly surface.On the initial
DO
part of the shell, it is substantially reinforced mid-
Kd
ventrally and lbrms a median ridge, similar to what we
| \,
know of other (fossil) sepioids.The delicatejuvenile shell, which has no solid rostrum to protect it, thus gainsin strength. (74) Sagittal grinding to the median plane reveals the internal structure(Fig. 28). The initial parts of the shell areparticularlyinteresting(Fig.2l). The septa are composedof mother of pearl and Fig. 28. - a. Median section through the last chambersof a
show a characteristicnacreouslustre. They are arched
Spirula shell, combining the data provided by Appellcif ( 1 8 9 3 a1 n dh i s f i g u r el . P I . 8 .1 2r n a t .s i z e .
in an hour-glass-likefashion, and close to the ventral
b. Section of a smal1piece of the outer shell wall. One can distinguish the three typical layers: hypostracum (fty), ostracum(os) and periostracum(po). (In the main figure, the first and secondlayers are combined).
side they grade into long calcareoustubes [septal necks].The latter continueeachas a horny (conchiolin) tube [the connectingring] which extendsbackwardsto the end of the precedingseptalneck, inside which it
x, y. anterior limit of mural part of septum; lrs. supporting
lies; at its end the connecting ring is united with the
ridge; sp1. last septum; sp2. penultimate septum; wk. living
surroundingseptalneck. The spacebetweenthem is
chamber;/k1.last gas chamber;/fr2.penultimategas chamber,
occupied by the pillar layer, which prevents the
forming only narow slits ventrally to the siphuncle(si); kd.
surfaces from being in contact and thus permits the
calcareousseptal neck; c/2. conchiolin connecting ring; pf.
passageof gases from the fleshy siphuncle to the
pillars; rd 1eftside ofshell aperture.
correspondingchamber.On the oppositeside (i.e. on the inside of the outer wall of the spiral shell) the septa are attachedwith a well-defined suture, although the
movementsthan on covering large distances.The loss
last layer of the fseptal]bowl gradesinto the innermost
of the rostrum coincideswith the animal's becoming
mother of pearl layer of the subsequentchamber.On
independentfrom the seabottom (p. 43) where sepioids
the posterior side the septal rim rests on a broad
typically live; a spike adaptedto burrowing would no
supportingledge which is deeply wedged betweenthe
longermake sense.
septumand the conotheca(cf. Parl I., p. l5).
A detailed considerationof the shell yields the
The first septumdiffers slightly in this respect:The
following points: 1. The protoconchis more markedly
constrictionfollowing the protoconchis so narrow that
globular and inflated (Fig. 27) than in other sepioids.2.
a normal developmentof the septumis impossible.The
The individual
chambers are.-.. demarcated by
septum therefore is rudimentary and immediately
c o n s t r i c t i o n s , t h e o n e b e t w e e n ( 73 ) t h e f i r s t
continuesinto the relatively large calcareoustube. The
[protoconch] and secondchambersbeing deeper than
latter of courselacks a point of fixation becausethere
others(Fig. 26c), the later onesbecoming increasingly
is no preceding tube; neverthelessits free rim grades
indistinct. 3. The last half whorl rapidly becomes
into a poorly calcified structure which here will be
detached from the earlier parts (so that the whorl
called the "conchiolin cap" (75) of the initial tube. lt
diameter increasesto nearly 6 mm), thus revealing
directly continues into the prosiphon, a poorly
similarity to typical sepioids(Figs 19, 23). Apparently
calcified. sasittal lamella of conchiolin which rests on
47 Fig.29. - Fossil shell ofSplrulirostrino lovisatoi Canavari. Nat. size.
From the Neogene near
Cagliari (Sardinia). ("Argille
fanghiane di
Fanganopresso Cagliari") (Miocene). Original specimenin the Stuttgafi museum. a. Dorsal view; preservedparts frced from matrix '' ii Si
in solid lines, reconstructedpafts in dotted 1ines. One recognizesa Sepia-likeshell. b. Lateral view, opened up to expose the
6 Vw
phragmocone and rostrum. Outline of the fposterior] end of the mantle sac added in dotted line. c. Ideal mediansection.Reconstructionfrom b. d. Ideal ventral view. Reconstruction from 6. using charactersin relatedforms (Fig. 23 and 30). Pr. pro-ostracum;l.s. last septum; Zw. ventral
,n !o
b.
wall (on the left at the free margin); -r. gas chamberin fork [not labelled:Vk?]; Sl. siphuncle; .l/r. lateral margin; Cp. capitulurn with protoconch;
-112.muscularmantle; Vf. ventralprocess;Do. spine (rostrum). [Vk, Vr not explained,probably meaning "ventral part of chamber" and "ventral margin", respectively].
the ventral-anteriorwall of the protoconch;in most
Here belongsonly:
cases its free (dorsal) edge is reinforced by a transversal lamella. The whole structure can be
The genusSpirulirostrina Canavari 1892.
morphologicallycompared to the pillars of the subsequenttubes, of which it is the precursor.The
With only one species:
secondseptumand its neck is alreadyformed nomally and shows all the parts distinctly. I am not certain whetherits conchiolintube is still closedposteriorly (asis its homolog in Nautilus),but I supposeit is.
Spirulirostfina lovisatoi Canavari 1892. This species is mentioned in textbooks (ZiItel, Grundztige 1921, p. 593), but it is not correctly describednor figured (the original descriptionwas not available to me, but I presume it was insufficient); I
I. The family Spirulirostrinidae Naef 192I
can now describe and figure it, with
(System,p. 536).
reconstruction,based on a specimenhoused in the
some
Stuttgart natural history collections. This is of some Diagnosis: Sepioideawith slender, delicate shells
importance since we see here a form apparently
intermediatebetweenSpirulirostra andSepia,having a
intermediatebetween Spiruliro,stra and the sepiids,a
well developed rostrum with a ventral process and
detail that was ignored (76) tn the descriptions
capitulum, which is neverthelesssmaller than in
availableto me2r.This can best be seenin Figure 29
Spirulirostra,
bands accompanying the phragmocone which in its
[Donovan: this reconstructionis said to be wrong by Janssen& Miiller 1984, Scripta Geol. 751 where
anterior part is stretched and dorso-ventrally
anyone familiar with the sepiids will be reminded of
compressed,the septa being ventrally drawn towards
slenderforms of cuttlebones(here Sepia fficinalis L.
the posteriorend so that they are convex and orientated obliquely, - in which the protoconchs are similar to
is of little use for comparison;but seeFig. 30e).
Spirulirostra, but slightly flatteneddorso-ventrallyand
viewed as a further preliminary stage leading to the
pushedinto an oblique position. (Other details can be
sepiid type, althoughthe age (Miocene) doesnot allow
takenfrom the reconstruction, Fig. 29).
(cf. p. 61) a direct placing into a palaeontological
in which the lateral edgesform narrow
In morphologicalterms the whole structurecan be
48
c.
d
Fig. 30. - The morphologicalderivationof the .Seplashell. a. A hypothetical intermediateform linking Sepia with Spirulirostra ("Spirulisepia", cf. Cephalopoda,vol. l, chapter40); this reconstructionwas made in 1914,before I was aware of Spirulirostrina (p. 76). While the stocky proportionsof this form do not conespondwith Spirulirostrina, the essentialorganisationis very close (Fig. 29), suggestingthat a rather slendershell forrrrwas also pdmary in sepiids. b. A typical shell of Sepla.Note (1) the replacementof the pro-osffacum(Pr1) by the lateralridges (fp) and the phragmocone(Is), (2) the degenerationof the ventralwall (Zu,) and the adjoiningpartsof the phragmoconewhich becomethe fork (GD). c. Belosepiasepioidea.A shelt in the Bavarian State Collections (public collections) in Munich. 112nat.size.Ventral view, very similar to Sepia.Thepart coffespondingto the ventralprocess(x) is serated,probablydue to macerationof the soft marginalpart. d. Ideal median sectionof c, with a carefully drawn alveolus.Phragmoconereconstructed(dotted lines). Note the archedprotoconch and the hom-like culature of the entire initial part. e. Laleralview ofthe shell ofajuvenile Sepia orbignyanaFdr.(cf. Fig. 32a).8,,nat. size. Due to the translucency ofthe delicate shell, one can seethe septalsutures,thus permitting comparisonwith Fig. 29b. The lateral ridges are similarly situated,facilitating the reconstructionof Spirulirostrina.F/. Wing-like parts of lateral ridges; Pr. pro-ostracum;,!p. lateral ridges; Is. last septum;Si. siphuncle;i'w. ventral wall; Cp. capitulum; Do. rostrum (spine);.r. ventral process(cf. Fig. 24. B);y. Belosepia;z. dorsalridge.
capitulum-like elevation in
49 Fig. 31. - Newly hatchedSepia fficinalis.aly nat. size. The dorsal view shows the animal in swimming position, at the very moment when the tentacularclubs are preparedto shoot out to catch a M1,sls.The tentacular stalks suddenly stretch out to catch the prey, which is caughtby the suckers,and are then rapidly retractedto deliver thc prey to the arms which hold it before the mouth, where it can be eaten. Note the overall similarity to Spirula (Fig. 26), very marked in some parts, whereas other parts are more typical of Sepla. The primary lid fold ofthe eye is closed,the slit-like pupil has a characteristicshape.The ventral anns have distinct swimming membranes, which - along with the slight ridges on thc ventro-lateralsides of the mantle (which becomesvery flat) - adhereto the substratewhen the animal restson the bottom. The outline of the fins shows the typical (often poorly seen) outline of the posterior end, whereas the anterior ends of the fins are still growing. On the dorsal side ofthe [posterior]end ofthe mantle,one can see an anchor-shapedglandulararea,"Hoyle's organ!'; it is involved in the hatching processand disappears soonaftcrwards.
"ancestralseries".
K. The family SepiidaeKeferstein1866.
The featureswhich support a close relationship with Spirulirostra are these: 1. The outline of the
Diagnosis: Sepioidea with a shell in which the
rostmml ventral processand capitulum.2. The distinct
periostracumis representedby an elongate-ovalto
ventral curvature of the posterior part of the
elongate-rhombic"dorsal shield" (79) in that the
phragmocone.3. The specialform of the initial and
anteriorly shifted "lateral edges" (p. 77) have almost
immediately following chambers.4. The secondary straighteningof the phragmocone.5) The outline of the
completely replaced the pro-ostracum,thus taking its topographicalposition, - in which the rostrurn in
lateralridges.All thesefeaturesare easily recognizable
generalis still representedby a powerful spine situated
when comparingFigures29 and23. (78) OIher featuressuggesta closer relationshipto
on the most posterior part of the shield, if it has not disappearedaltogether,- in which the phragmoconeis
sepiids: 1. The initial part of the phragmoconeis
divided into a foliaceously chambered "hump" that
already compresseddorso-ventrally,and the initial
reachesnearly to the anterior end of the shield, and a
chamber is not globular or bubble-shaped.2. The
lower part called the "fork", lying ventrally to the
rostrum is comparativelysmall, as in Sepia.3. The
siphuncle;both parts show very oblique septaslanting
later chambers appear increasingly flattened and the
down from the antero-dorsalto the postero-ventralend, - in which the siphuncle widens very rapidly thus in
septaare not perpendicularto the phragmoconesurface but strikingly oblique and the suture, also in lateral aspect(Fig._29b),curved posteriorly.- I presumethat
parl forming a mere depressionrather than a narrow
theseundeniablesimilaritiesto sepiidsare linked with
tube; during later developmentthe depressioncan even becomeconvex"- in which the ventral arms now have
othersthat are not yet recognized,in particular I think
two very powerful, fin-like swimming membranes,and
that the siphuncleis funnel-shapedanteriorly and that
the fins grow far forwards on either side of the mantle
the ventral wall is reduced.The resulting form would
sac.(SeeCephalopoda, vol. I, chapter40).
be the one given in the reconstruction(Fig. 29c, d).
The anatomy and externalmorphology (Fig. 3l) of
50
Fig, 32.
Median sectionsthroughthe mantlesacand shell of sepiids,slightly schematic.
a. Shell of Belosepiasepioidea,like Figs 30 and 33 (nat. size). b. Sepia orbignyana (uvenile), a form rather close to typical Sepiinae.Note the strong ventral curvature,the embedding of the phragmoconein the sheath(the latter is marked black, as in a), the arangement of the pillars and intermediatesepta (Zs), and considerthe greatextcnt ofthe siphuncularpit and the pafts coffespondingto the septalnecks.18/1nat. size.1. c. Sepia o.fficinalis.One recognizesa simplified form of rostnrm and a modified fork and ventral wall (Ga). The latter has grown normally for some time, but then the conothecacurved outwards and the fork lamellae continued to be depositcdon the ventral processtUg). t: I nat.size). d. Mantle sac in median section.The muscularmantle(Mm) insefis on the traditionalsite (Fig. 248), i.e. on the ventralprocess(tr/r. (Nat. size).
the soft body cannotbe discussedin detail here,but the
phragmocone.The latter, followed by the lateralridges,
architectureof the cuttlebone deservesour special
has becomeextendedto the anterior end of the mantle
attention (for which we have been prepared by the
sac (in fact reachingbeyond the nuchal attachment),so
foregoingchapters):
that the septa had to assumea very oblique position ( F i g . 3 0 ) . D u r i n g t h i s p r o g r e s s i o inn a n a n t e r i o r
First of all we have to emphasizethe total loss of an integrantparl of the typical dibranchiateshell, namely
direction, evidently the ventral wall did not keep pace.
the pro-ostracum;this loss may suggest close
If it had, it would have penetratedeven more deeply
relationshipto Spirula (p. 72).A closerlook, however,
into the soft body than the actual situation in typical
indicatesthat this part in Spirula is replacedby the
sepioids (cf. p. 33 and p. 47). Instead,it has been
muscularmantle,whereasin Sepia it is replacedby the
reduced in a striking fashion in all sepiids; it thus
51 appearsrestrictedto the most posteriorparl of the body
features (a distinct ventral curvature of the strongly
in the sepiids,especiallyin the recentforms (Fig. 35).
arched first chamber, a narrow siphuncle, widely
The siphuncle,which was probably alreadyextendedin
spacedsepta), in which the ventral wall of the shell
Spirulirostra, is enormouslystretchedand thus widely
(fork) preservesa structuresimilar to Spirulirostra, in
separatesthe ventral and dorsal parts of the septa.A
which the gas chambersof the "hump" are not fully
result of this separation(80) must have been the structuraldifferenceof the septalcomponents,rvhich is
equipped with a scaffolding made of pillars and intermediate septa, - in which a well-developed
well known in the modern cuttlefish; it separatesthe
rostrum,which is dorsally curved,is alwayspresent,its
posterior, ventral part called the "fork" from the anterior,dorsal part called the "hump" (Fig. 32b). The
base continuing medio-dorsallyinto a warty, crest-like differentiationof the sheath,- in which the ventral side
septa of the fork are connected laterally with their
of the most posteriorpart of the dorsal shield shows a
countelpaftsin the hump (Fig. 30b).
sculpturewith distinct grooves radiating out from the
Moreover, the angle of the aperture of the phragmoconeincreasesgreatly, as is recognizable
baseofthe fork (Fig. 30c,d, 32a,33). (82) Here belongs:
already in the initial chamber. (81) Instead of being bubble-shapedit is rather flat, bowl-shaped.This
The genusBelosepiaVoltz 1830.
entailsthe disappearance of a typical capitulum,which is totally absentin the sepiidsproper(Fig. 32). Finally, the componentsof the sheathare modified in both their positions and proportions:The lateral
with
numerous species, some insufficiently
characterized,the distinction of which is of no special interesthere.Theseare:
plates have followed the phragmoconeto the anterior end of the mantle sac, where they unite with an oval
Saepiat Cuvier 1824(Annales),Pl. 22, Figs l-3.
outline, since a free pro-ostracumis totally lacking. In
Sepia cuvieri d'Orb. 1825,p. 67, Deshayes1825. P1. 101,
the opposite,posteriordirection they reach,in a similar fashion, the midline where they becomeunited via an intermediatepart, which is ontogeneticallydistinct and
Figs 7-9. BelopterosepioideaBlainv. 1825,p.622,Pl.59, Fig. l. S e p i a c u v i e r i S o w e r b y1 8 2 9 ,P I . 5 9 , F i g . 1 . H e r e a l s o . l . brevispino.
which correspondsto the processus ventralis (p. 6l).
Beloptera cuvieriBrckland 1836,Pl.44', Fig. 15.
The greater part of the peripheral sections of the
Belosepiacuvieri Yoltz 1836,p. 6.
sheath,on which the muscular mantle is inserted,has
Sepia longispina,longirostris, blainvillei Deshayes1837, Pl.
thus becomethe continuousrim of the "dorsal shield"; the periostracumnow appearsin the form of a flat to more or less strongly archedplate accommodatingthe phragmocone(Fig. 32b). At the posteriorend of this plate, the rostrum is representedby a (relatively) inconspicuousspine which no longer envelopesa
1 0 1 p, . 7 5 7 . SepiactrvieriQuenst.1849. Belosepia .sepioideaEdwards and Wood 1849, 1877. Sepiasepioidead'Orb. I 850, Prodr.2, p. 338. Here also S. cuvieri, longispina, longirostris, blainvillei Desh. BelosepialongispinaChenu 1859.p. 46, Figs 140, 141.
considerable part of the chambered shell. More
BelosepiasepioideaChenu 1859,p. 46,Fig. 142.
anteriorly crest-like elevations,often in a mid-dorsal
Belosepiasepioideavar. longirostris,p. 46, Figs 143, 144.
position,appearon the "dorsal shield".
BelosepiasepioideaKeferstein1866,p. 1441,Pl. 120, Fig.
Considerabledifferencesin detail appearbetween the Belosepiinaeof the Eocene (Paleocene)and the Eusepiinae [?Sepiinae] which appear only in the Miocene.
\l-t2 BelosepiasepioideaFischer 1887,p. 557, Fig. 135 (drawing after Mun.-Chalmas) Belosepiablainvillei Zittel 1885,p. 514, Figs 707-708 BelosepiasepioideaCrick 1894,Pl. 10. BelosepiablainvilleiPompeckyl9l2,p. 1961,Fig.58. Sepia vera Desh. 1866, Pl. 106, Fig. I l-12. Cossmann&
I . The subfamilyBelosepiinaeNaef 1921.
P i s s a r o1 9 1 3 P , I . 6 0 ,F i g . 1 - 1 . BelosepiasepioideaBlainv.. Cossmann& Pissaro1913.Pl.
Diagnosis: Sepiidaein which the phragmoconeretains, at least in its earliest part, rather typical sepioid
60, Fig. 2- I .
52
A,
r' ,[
Fig. 33. - Shellsof Belosepia. a. Dorsalview. D. Median sectionof a shell of B. sepioideaafter E,dwardsand Wood (1877), almost completelypreserwed.r/2 nat. size. Note the curvature"the courseofthe sufures,the ventralwall. c. A fragmentfrom the public collectionsin Munich (BavarianStateCollections),which is shown in a (constructed)median section ind. e. Ventral view of the posteriorpart of c. /. Dorsal view of a fragmentin the British Museum,after Crick (1894,Pl. 10). c1. Posteriorend of the alveoluswith septa.septalnecks, pillars, suture lines etc., drawn from Edwards (1877, Pl. I, Fig. 6) (interyretedand improved) and combined with c so that it lies in the natural position. .s.last septum,followed by two more suture lines.
Belosepia ov,eniDesh.,ibidem,Ftg. 2-2. Belosepia brevispina Sow.,ibidem,Fig.2-3. Belosepia blainvilleiDesh.,ibidem,Fig.2-4. Belosepia tricarinataWatelet,ibidem,Fig.2-6. (Stenosepia) B. compressa Blainv.,ibidem,Fig.2-5. The morphologic-systematic position of these fossils was alreadycorrectly determinedby Cuvier (1824), as indicatedby the namewhich he used.Blainville(1825) recognizedthe relationshipto more typical sepioids (our Beloptera) and Voltz (1836) appreciatedits intermediatecharacter.Buckland (1836, Pl. 44', Fig. 15) considered"Beloptera" as an "intermediate form between a belemnite and the shell or sheathof Sepla
shells are strongly curved in both longitudinal and transversesection,so that the shapeof the animal must have been laterally compressed,in contrastto the more or less dorso-ventrallv flattened shaoe of the Eusepiinae. In this respect,they were apparentlystill closer to the other sepioids,so the special shapeof the ventral arms in Sepia, which is related to the shapeof the mantle (Fig. 31), cannot necessarilybe considereda featurealreadypresentin the Belosepiinae.In the latter the extent of the fins was probably limited to the posterior end. But in other respectstheir organisation must havebeensimilar to that of Sepia.
o//icinalis", which he illustratedvery well (loc. cit., Fig. a). This partly inappropriateinterpretationwas adoptedby more recentauthors(e.g. Lang 1900,p. 99,
(84) 2.The subfamilySepiinaeNaef 1921.
Fig. 107b). (83)
Apart from many vague similarities to
Diagnosis: Sepiidswith a phragmoconeshowing very
Spirtrl irostra and Spirulirostrina the Belosepiinae
early the typical featuresof the family, namely a flat
exhibit several essentialpecularitiesof their own: the
initial chamber,a flat and broad siphuncle,closely
53 spacedsepta with a distinct separationof the parts belonging to the hump and fork, respectively.- in u,hichthe gas chambersdo not typically extendinto the fbrk (ventral wa11), in which the flat chambersare equippedwith a scaffoldingof pillars and interrnediate septa,- in which the small or missing rostrum is very rarely accompaniedor even replacedby a small, knotty longitudinalridge. Here belongs
in addition to some rare modern
types that are of no special interest in the present context2r- : Tl.regenusSepiaL. This genuscontainsa large number of living and about a dozen fossil species.The latter are more or less well defined; they are known from the Miocene onwards
Fig. 34.
and in
the posteriorsidc. 1. positionof the reducedcapitulum(Fig.
every respect resemble the
living
representatives, which will thereforebe treatedfirst:
The fragment shown in Figs 30. 33 c-rd seenfrom
32a); 2. remains of the ventral wall; 3. inner surfaceof the latter,with fork 1amel1ae; 4. courseof the fork; 5. first suture
a) The cuttlebonesofRecent Sepia. The general outline of the cuttlebone ("Sepion" "Sepiostaire","Osseletinterne") is well known from
(protoconch);6-8. further sutures; 9. fracture surface; 10. cristal 11. rostrum(nat. size).
lield collections and from illustrations in handbooks and textbooks,as is the aspectof the major parls: 1. the
(seeabovep.76-78). discussed
dorsal "shield" ("bouclier", "Riickenplatte"),2. the "rostrum" ("Dorn", "spine"), 3. the "hump"
speciesand a relatively high diversity of forms, so that
("Spongioidtissue"), 4. the "fork". Voltz (1830)
data for historic-morphologicalstudiescould hopefully
already knew that the last two parts correspondwith
be extracted from it. Unfortunately the majority of
The genus Sepia now shows a great number of
the phragmocone,and he aiso recognizedthat the
speciesis poorly known, however,and in most cases
innermost layer of the shield ("teste alvdolaire")
the juvenile shell is virtually unknown. We therefore
corespondsto the conothecaof a belemnite,while the
limit our present considerationsto the shell of Sepia
rest must be consideredas a sheath("gaine"). Riefstahl
ofJicinalis,which is the most easily availableform, and
(1885) presentedthesehomologiesin greaterdetaiPs.
then attempt to embark on some comparative
A more detailedanalysishas only been given by (85)
considerations.
Appellof (1893)who studiedthe microscopicstructure and its developmentin thin sections. Little can be added to the descriptionsgiven by
A generaloverview of the shapeand position of the shell in an advancedembryo is given in Fig. 8d, which will be our starting point. lt broadly shows the same
Appellof; it will be included in the forthcoming
conditions as in the adult animal, but clearly the
description of cuttlefish from the Naples area
curvatureis much more pronouncedand the chambers
(Cephalopoda,voi. I, chapter 40). What appears
are more distinctly visible. (The skin and the shell are
essentialhere is the relation to a morphological series
delicate and translucent!). The excentric growth
of which Sepia is the ultimate member.This leadsto a
reminds one of the typical ventral curvature, and the
more detailed and instructive picture of the homology
individual septawhen seenfrom below (Cephalopoda,
of the individual parts and conditions, as well as
vol. II, Pl. 20, Figs 5, 6) initially clearly show circular
providing a rational context. For in the serial
depressionsinstead of a siphuncle.In the first few
arrangementeachmemberprovidesthe prerequisitefor
septa,differentiation of hump and fork parts (cf. Fig.
u n d e r s t a n d i n gt h e s u b s e q u e n to n e s , a s a l r e a d y
38) is not yet visible; (86) lt appearsprogressively
54 Fig. 35.
b
Moryhology and topographyof the shell of
Sepio fficinalis. a. She1lof a maturing female.
3/5 nat. size. This shell
belongs to a variety in which the siphuncularpart is very short and remains limited to the posterior half of the body, in contrastto .'. b. Shel1of a juvenile individual, in which the fork still shows normal structure,in that the ventral wall (i/w) projectsfreely. embeddedin the soft tissues(Fig. 32a). 3/2 nat. size. C/r. uncalcified, chitinous shell margin; Kr. calcified part of the she1lmargin, appearingwhite; Sx'. last f.i,
septum of the hump; Si. siphuncular pafi: Ga. fork; Lg upturned wall of the fork, diffcrentiatedfollowing the juvcnile stageofthe shell, in which the free edgeofthe
.
conotheca (Rd) and the vcntral wall (l/u,) were stil1 visible. c. Shell insertedinto the soft body, showing especially the relation to the muscular mantle (M), thc primary mantle (N/r) and the head-footretractors(t-1).315 nat. size."Emptied mantle sac". The shell is visible through thc siphuncularepithelium.The anteriorpart. abovethe cur-vedline betweenthe stellateganglia (which marks the posterior limit of the dorsal mantle cavity), is only covered by the primary mantle, which here has a cartilaginous structure ("nuchal attachment") and normally adheresto the "co1lar attachment"(Fig. 3b). Medially to the stellategangliaone can seethe cut ends of the large muscleswhich inserl on the free margin of the shell and on the lateralparls of the hump, in fact on the she1lsac which covers thesc parts: the head-foot retractors (1, 2, 3) and the funnel retractors (4). The posterior parts of the lateral plates are covered by the muscularmantle,which here insertson the ventral side. The fork, the median parts of the siphuncularpit. part of the last hump lamella and two small comers of the shield margin (x) are covered only by the shell sac. without muscle insertions. All the viscera have been removed,the branchialbands (Kb) severedclose to the mantle.
distinct only by the time of hatching,which occurs
later becomessplit up fto form intermediatelamellae]. However, the pillars (Fig. 37) are present from the
after formation of the eighth septum. (Premature
beginning. In the earliestchambersthey form isolated
hatching is common under aquarium conditionsl).
beams,which only later becomeunited as longitudinal
Embryonic shells already lack a pro-ostracum;
septa running paralle1 to one another ("vertical
however, the (soft) mid-dorsal margin (87) of the
lamellae"). The lateral platestaper posteriorly (cf. Fig.
dorsal shield is slightly lengthenedand pointed in the
38) and are united by a solid, curvedrim (Fig. 37a:20),
early stages(Fig. 38f, g). The last-formedchamberis
the ventralprocess. which doubtlessrepresents
during subsequentgrowth stages.The rostrumbecomes
always filled with gelatinousconchiolin,which only
Starting from these embryonic conditions,those of
55 Fig. 36.
Typical crosssectionsofthe posterior
part of the body and shell of .lepla ofticinalis, to illustratethe topographicalrelationsbetweenthe muscular mantle. fins, head-foot and funnel refraat6rc
oillq
cfc
a. Most posteriorpart (i.e. zone of the siphuncle and wings). D. In the middle parl of the shell (anteriorto the siphuncularpart). CompareFig. 35 and note that in a the muscular mantle inscrts on the ventral side. in D on the (bent down) dorsal side of the shell. The fins lic. via an artioulatedconnection, on the most posteriorpart ofthe shell and on the muscular mantle (c1-.above p. 34, Fig. 71). The powerful head-foot and funnel retractorsinsert on the inner surfaceofthe lateralplates.anterior to the fork (Fig. 35c); they partly encroachon the hump and flank the liver. (cf. Spirulo.Fig. 26c). KD. branchialband; Ka. branchialartery;Kv. branchial vein; Kru. branchial splecn; 7g. ink duct; fc. Vena cava; Ed. intestine; F/. fin musculature; Ft.
Iln cartilage; I,l/t. fin-base
pouch (cf. p. 34); Scft. shell (lateral plate); -r-r. siphuncle;Ga. fork; S.r.shell sac; Co. coelom; Mh. mantle cavity; Ms. mantle septum: Mrr. muscular mantle; Rd. bent down shell margin; Sp. septum; iila. hump; Mlr. muscular mass (1: lread-footretractor,2: funnel rctractor);B/r. attachmentfold of gill lamellaB/; Lb.liver. c'. Shell between a and b, without the soft parts. In the anterior part ofthe oldest chamber (Ift), one can recognize the niches mentionedabove(p. 87); in the subsequent chambersthe intermediatesepta(In) arevisible, and in all chamberstherearepillars (Sl). Sp. lateral plate; Go. fork; Co. conotheca;Sl. pillars;1n. intermediateseptatSu'2.the secondscptum sectioned;2ft.oldest chamber; Rp. dorsal shield (periostracum);-r-.r. siphuncularpart; Sl. Septal neck; Sw7.convex surface of siphuncle,i.e. the earliest neighbouringscpta.cut at the transitionto the scptalneck (cf. Fig. 30b).
the juvenile shell (Fig. 35b) becomeimmediately
other interesting aspects:first of all it should be
understandable. Noteworthy are the posterior tapering
emphasizedthat the supporting lamellae (pillars)
of the lateral edges, and the rather narrow ventral
become more regular anteriorly, lying ahnost parallel
process,which largely revealsthe small spine. It is
to one anotherand progressivelydividing along their
particularly characteristicthat the ventral wall is still
length to finally diverge to some extent in a fan-like
presentin its typical, upright position, whereaslater on
pattern.At the anterior end of each chamberseparate,
it will be overgrownby the septaof the fork (Fig. 32).
narrow spaces are thus formed (separatedby the
The juvenile shell of Sepia fficinalrs (and many
"cloisons verticales").The intermediateseptado not
other species)thus resemblesthe cuttlebone of Sepia
extend that far. A11supportinglamellae have a nearly
acLtleatad'Orb. and its allies, which never develop
smooth rim where they rest on the previous septum,a
beyond that stage. (This group is characterizedby a p o o r l y d e v e l o p e dp o s t e r i o r m a r g i n , w h i c h i s n o t
peculiar meanderingrim at their insertion on the new
strongly curved,by a free spine,and by an upright fork
suturelines of ammoniteshells,and we may indeed
in the posteriorparl of the shellza).
interpretthe similarity of this structureas the result of a
The detailed structureof the humo shows some
septum.This results in a pattern reminiscentof the
similar bio-mechanicalcause:apparentlythe prop is
56 thus more evenly distributed on the surface needing
freely into the soft body, and the lateral edgesprovide
support,with the additionaleffect of greaterelasticity.
the insertion points for the head-foot retractors, the
(88/ This indeedwas the interpretationof Buckland
homologouspoints in Sepia were still obscure:we
(1836) ( cf. Agassiz1838)who remarked(explanation
finally found them at the edge of the fork where the
to Pl. 44'): "The rneanderingform of thesewalls has
correspondingmuscles indeed extend backwardsto
the effect that they, like the leaf-shapededges of
insert. Anteriorly a shift occurred due to the oblique,
ammonitesepta,resistmore readily to pressure".
(90) nearly horizontal positions of the septa.But the
Detailed considerationsof the insertion of the soft
typical similarity is obvious when comparing Figures
body in the muscularmass are necessary(Fig. 35c).
26 and 35[c], including the way in which the funnel
There is no further need to refute a simple statement like "the shell is stuck in the mantle". (89) lt is clear
retractorsjoin the head-footretractorslaterally. Thus a
that in a sepiid the origin of the muscular mantle no
intermediateforms, which could even be useful to
longer coincideswith the primary shell margin (cf. p.
peoplenot immediatelyinvolved in this study.
81 and Fig. 30). We indeed see a "secondaryshell
solid basis is available for reconstruction of
Some architecturalconditionsof the shell are given
margin", which is largely homologouswith the lateral
in Fig. 37 using schematic median sections: this
edges which now extend to the anterior end. At the
particularly applies to the layers of the dorsal shield,
posterior end, however, the morphological equivalent
which has beenconectly describedby Appellof ( 1893)
of the "ventral process" adds a special feature to the
but which had never and by some of his predecessors,
relationshipbetweenthe muscularmantle and the shell.
been fully explainedin morphologicalterms.We can
The lateral plates also deviate somewhat,as seems
of course expect to find three main layers: the
understandablewhen looking at Spirulirostra: in the
ostracum,the hypostracumand the periostracum.The
posterior part, which is somewhatlaterally expanded
former two are easily identified when starling from the
(more strongly in some speciesthan in the present
chamberedpart of the shell (Fig. 37c: 5 and 6). If we
one), thus forming the so-called"wings", the muscular
start from the dorsal side, however, we find complex
mantle insertsas in the ventral process,i.e. on the side
and confusingconditions.The whole marginal zone, to
of the secondary shell margin facing forward and
the line 3, is uncalcified,flexible and could easily be
downward (cf. Fig. 2482), thus reaching almost to the
c o n s i d e r e da s a s p e c i a l m o r p h o l o g i c a lu n i t . B u t
fork (Fig. 35c). At the point markedx fon Fig. 35c] it moves outwardsagain to occupy the free (uncalcified)
apparentlythis is only unfinishedmaterial,since the
rim. This rim here is curved in a peculiar way (Fig.
parls ofthe shell. The subsequentlayers(1, 2) are very
36b: Rd) so that the muscularmantle now reachesthe
distinct, the lower layer being the thickest one in the
outside of the dorsal shield and thence extendsto the
anterior part of the shell. If the outer layer only were
anterior margin; here the curved rirn is somewhat
consideredto representthe sheath,as seemsevident,
extended and the muscular mantle is reduced to its
there would be no convincing interpretation for the
smallestextent,reachingthe oppositeside only as a
inner layer. Once we have recognizedthe true natureof
narrow band. This arrangementis inherited frorn the
the lateralplatesand their extensionto the anteriorend
pro-ostracum,which has been replaced by the lateral
in sepiids, however, we have the solution of the
edges,whereasthe arrangementof the wings strikingly resemblesthat of the conusvane in Trachyrctnhis(Fig.
problem: the sheathfotms a free edge and reachesalso
52), thus providing a beautiful example of atypical
accretionon both the upper and the lower sides.On the
similarity ("convergence"). Indeed, a
close
latter (7-8) the matrix forms a special plate
The insertion of the head retractorson the shell is
p o s i t i o n a n d p r o g r e s s i v et h i c k e n i n g . T h i s p l a t e
also notewofthy for comparisonwith Spirula (Fig. 26).
continues (along with the corresponding strip of
The transitionalstagesof Figs 23,29 and 30 will have
thickened shell epithelium) directly into the lateral
to be taken into considerationto achieve a new
edges; we thus have an interpretation which fits the
illustrationof the unity of type in the sepioids.Whereas
factsperfectly.
phylogeneticrelationshipis out ofthe question.
rn Spirula the primary shell aperlureclearly penetrates
samelayerscan be tracedas calcified ones in the older
anteriorly beyond the phragmocone; it grows by
correspondingto the hypostracumin terms of both
Interpretationof the phragmoconein the areaof the
)/ Fig. 37. - Median sectionsof the shell of Sepia ollicinalis (in parl after Appellcif 1893.interpretedand simplilied). a. Embryonic shcll, after formation of the first septum with siphuncularpit (18). For a better understanding,the primary shell (21, ostracurn and hypostracurn) and the secondary shel1 (22, periostracum)are distinguishedhere, althoughthesecannotyet bc distinguishedin prcparations.either in dccalcified sectionsor in sectionsof the shell (cf. Appelldl Pl. 6, Fig. 2). The flattened initial chamber appearsembeddedin the sheath whrch already showsthe reflectedventralprocess(20). 19. pillar in the region of the initial sipl.runcle, correspondingto a prosiphon(Fig. 21). D. Section through the pafi close to the siphuncularho11ow(cf. Appellof,P1.2, Fig. I andPl.5, Fig.4). 1. septum;2. septalneck of the fbllowing septum;3. "deposit in the posteriorcorner of the chamber"; 1. gas chamber; 5. pillar of gas chamber; 6. intemediateseptum(cf. Fig. 32 at Sk). c'.Median sectionof the anteriorend of the shcll (cf. Fig. 32a and Appelldf,Pl. 2, Fig. 2). 1. outerlayer ofperiostracum(Appellof's "dorsal plate"); 2. inner layer of the latter ("median plate"); 3. limit of calcification; 4. calcified part of median plate; 5. ostracum,6. hypostracum("black. stratihedlayer ofinner platc"); 7. growing anterior margin. belonging to thc periostracum; 7-8. glowth zone of inner layer of periostracum;B-9. growth zone of the ostracumi 9-10. growth zone of the hypostracum(a special pafi of the shell cpithelium. providing the formational matrix, correspondsto cach of these zones)l I1-16. soft, gelatinous, stratilied shcl1 material with pillars (1 Z) incorporated in it, a prospective"cavity layer" which will form cavities (due to shrinkageof the fi1ling) only after the formation of the following septum; 12. insertion of the preceding septum (15) on the hypostracum;13. supporlingridge ("deposit in the anteriorcorner ofthe chamber");11. pillar; 15. (see12!); 16. (see1 1t):17. (see l ll); 17a. intermediateseptum;/c.last chamber.
hump raisesno difficulties. One observesa continuous
the chamber(Fig. 36c) but are regularly formed in the
s e c r e t i o no f s o f t c o n c h i o l i n ( F i g . 3 7 c : 1 6 ) w i t h
posteriorpart (Fig. 36b). Supportingridges (Fig. 37c:
integrated pillars (17), which alternates with the
13) are alsotypical.
formation of septa(15). The resultingchambers(k) are
(92) The ventral side of the hump (Fig. 37b) shows
initially filled with conchiolin, which only later
the last septum and the strip-like homologues of the
contractsand is replacedby gas (91) (Appelldf 1893,
septal necks (2) of all the preceding septa (l). A
Pl. 5). The shrunkenconchiolin lorms the intermediate
peculiarity of this part is a modificationof the pillars in
septa(17a), which are absentfrom the anterior part of
the septal neck zone to form a sort ofjoint (3) in the
58 median part. Apparently pillars are lacking in the
I have no definite opinion abottt Plagioteuthis
lateralparts (cf. Appellof 1893,Pl. 5, Figs 1 and 3);
moscoviensis (Rcimerand Damas 1890). There is a
they can be interpretedas a modification of the pillars
ceftain similarity to Belemnosis(cf. Fig. 15). But again
in the "posteriorchambercomer".(cf.Fig.27a).
a more detailed study is necessary.It is out of the questionto basethe assumptionof a greaterage of the
b) The fossil speciesof Sepia.
whole suborder on this incompletely preserved
Numerousspeciesof fossil cuttlefish are known, based
fragmentostensiblyfrom the Lower Malm (Oxfordian)
solely on cuttlebones,which are often more poorly
of Russia.
characterized(p. 8a) than the shellsof living Sepla.As
As regardsa third form that has beenplacedhere,it
far as I can gather from my own observationsand from
is even more certain that it does not represent a
data in the literature,all of these forms are very close
cuttlefish-like animal. This is "Campl,losepia" (Picard
to the living forms and thus cannot tell us anything
1899).Two specieshavebeendescribed:7. C. triassica
about the earlier history of recent cuttlefish. In some
P i c a r d 1 8 9 9 ,p . 3 0 8 , P l . 1 a n d F i g s 1 3 - 1 4 ,a n d 2 . C .
specimens,especiallyin those from the Miocene marls
elongataPicard 1910,p. 359, Fig. 2. Anyone familiar
of mainland Italy and Sardinia,one hnds (as in some
with sepioid shells will find no reasonto acceptthese
recentspeciesof the aculeatatype) a slight protrusion
horn-shaped fossils in the present context. As a
of the rounded anterior margin, which may suggesta
geologist,the author of these speciesmay have seen
vestige of an earlier pro-ostracum(cf. Fig. 38). We
some analogy to the rostrum of cuttlefish. The only
have no fossilsof the genusfrom earlierstrata25. The
noteworthy detail is the geological age of this
Sepiinae from the Miocene were apparently derived
ostensiblesepiid, which originally came to me as a
from belosepiidsof the Eocene,from which they differ
surprise.Indeed, since the typical predecessorsof the
in addition to the morphologicalfeaturesalready
sepiids are not known from strata older than Eocene
by the delicate nature of their shell and
(Paleocene),the occurrence of derived forms in
their smallersize.Their size corresponds to that of the
Triassic strata would upset our understandingof the
smallerrecentspecies(e.g.S.orbignyana).
group.
mentioned
Examplesare: L S. michellottii (Gastaldi1868,p.
The same can be said about another "cuttlebone",
226, PI. 5; a complete shell from the marls near
allegedly from the Devonian, named "Palaeotettthis
Superga,Middle Miocene).2. S. craveri (ibid. Pl. 4)
dunensis"Romer26(cf. Rcimer 1856,p. 72, Pl. 13).
from Pliocene marls of Brd. 3. S. vindobonensis
There is some doubt whether the material in which the
( S c h l o e n b a c h1 8 6 9 , a c o m p l e t e s h e l 1 ) f r o m t h e
fossil was found embeddedis really a greywacke of
Neogene near Vienna (Tegel von Baden). 4. S.
Devonian age, since only castsof dissolvedshells(94)
lovisatoiParonaand 5. S. calaritana Parona,both from
Accordingto Zittel (l 885,p. could be expectedthere27.
Miocene marls near Cagliari, Sardinia (Fargeri,
521) thesestructuresareplacoganoidscales!
Fangario).Sacco(Bellardi)reports(1904,p. 3-4) on 10
Sepion (?) taurinense Sacco can hardly have
speciesfrom the Miocene of Italy. (Also seev. Biilow,
anything to do with sepiids;nothing in the problematic
1921, p. 247-248; S. sttbsagittata belongs to
fragment speaksfor such an association.The author of
Plesiotufihisl).
this speciesplacesit only tentativelywith cephalopods.
(91 Some reports on cuttlebone-like shells from earlier periods are likely to generatemore confusion;
Likewise Orcagnia Oppenheirn1899doesnot belong here.
the following sectionis thereforeadded: Appendix: Spurious sepioidsfrom older strata. Here I discussa problematicform that cannotbe a
L. Review of the fossil Sepioidea and their evolution.
" s p i r u l i d " ( i . e . a s e p i o i d i n o u r s e n s e ) ,n a r n e l y "Atrloceras" (Trautschold1866). The phragmocones
In summary we can state that it is possibleto arrange
(providedthey really were phragmocones)are from the Lias of Russia; their detailed assessmentrequires
the decapodtypes mentioned either directly or by subjoining them - in a morphological series
fuither study(cf. Biilow 1920).
intermediatebetween the belemnitesand the recent
59 sepiids.This is especiallythe casewith the following
2l), alw,ayspermit a certain ecological interpretation.
forms:
We have to keep this constantlyin mind; such a view B e l e m n o s e l l a a m e r i c a n a ,2 . S p i r u l i r o s t r a
which is no longer morphologicallybiasedleadsto the
LteI I ar d i i, 3. Sp.sepioidea, 4. Spirtrliro strin a lovisa toi,
1.
impressionthat the real developmentdeducedfrom the
5. Belosepiasepioidea,6.Sepiaspec.(Miocenespecies
systematic-morphological seriesreflectsa continuous
and the groupof S. aaileata),7 . Sepiafficinalis.
processofecological adaptationin certainspecies:
This seriescould possiblybe extendedbackwards
l. The modification of the belemnoid type to
(Fig. 6d) to include Diploconus belemnitoide,.r or a
Belemnosella
similar belemnoid form with a bulky rostrum; it first
adaptations:l. The strengtheningofthe activeparts of
leadsto a sepiidtype best represented by Sepiarostrata
the organisation as opposed to the passive ones
apparently resulted in different
or S. actrleata d'Orb. D'Orbigny (1842, Annales,p.
(muscularmantle versusshell), a phenomenonthat can
363-365)alreadyrealizedthe close connectionbetween
be observedin most cephalopodlineages(cf. Fig. 39).
numbers 5 and 2 of our series.This fact should have
2. The stabilisation of equilibrium by shifting the part
led much earlier to the establishmentof my suborder
of the shell containinggas to the dorsal side (p. 47), as
S e p i o i d e a .I n d e e d , d ' O r b i g n y c o n s i d e r e dS p i r u l a
can also be observedin other examples(endoceratids,
rvithin this family relationship2s.
ascoceratids). 3. Through the advantagesthus obtained,
These highly interesting relationships were
Ihepossibility (free energy)arisesto achievedivergent
subsequentlyneglectedin zoology and palaeontology.
modi.fications and progressiveexploitation of new
Thus, Zittel (1885) simply included Belenmosis,
living conditions, as reflected by the diversity of
Beloptera,Belopterina,Spirtrlirostrain the belemnites,
sepioidforms (cf. p. 24).
riith which he associatedSpirula at subfamily level.
(96) Il. Compared to the preceding one, the
The "Sepiophora" (Teuthoideaand Sepiidae)are then
Spirulirostra type is again characterizedby a number
s h a r p l y s e p a r a t e d .T h e f a m o u s " m o r p h o l o g i c a l
of advantages:the broadening of the lateral bulges to
primacy", which is supposedto recognize systematic
wings (as rn Beloptera) results in more space for
relationshipsamong different forms, (95) here clearly
visceraldevelopment,which is also enhancedby the
f-ailedto generatea brilliant result! It shows how
ventrai curvature of the phragmocone.This offers the
necessary this revisionin fact is.
possibilityof an increasein bod.vsre; this possibility
It cannot be denied that in certain particularsa
was remarkablyexploitedby Sp.hoerneslin which the
different opinion is possible. Before I knew of Belemnosella antericana, I indeed considered
shell adopted a particularly broad arch, thus a p p r o a c h i n gt h e s h a p e o f s e p i i d s h e l l s . - T h e
B e l e m n o s i s c o . s s m a n na i s the initial form of the
improvement of ntuscular mantle insertion was
sepioids(p. 49). Starting from this form, one could easilyget - via Spirulirostridittntobtusum(p. 62.y- to
probably relatedto an increasein mantle efficiency, againenhancedby the shapeofthe posteriorend.
Spirtrlirostra bellardii and then proceed to the same
IlL ln Spirulirostrina the muscular mantle is
derivation of the sepiid type. That Belemnosella is
becoming more independentof the shell: its shapeand
rnore likely the ancestralform is suggestedby its close
size appearsnot so much determinedby the shell as in
relationshipto the belemnoid type, i.e. to the general
Spirtrlirostra (Fig. 2q, not even at the posterior end.
type of decapods,which has the "morphological
There is virtually no (architectural) impediment to
primacy" (Naef 1919, p. 29). Belemnosisand
enlargementor broadening,to flatteningor rounding of
Spirulirostridium then appear as "secondary forms"
the mantle sac, so that the way to modification towards
characterized by an obtuserostrum.
the sepiid type is open. The initial shift of the parts (p.
In my view, Spirulirostra appearsas a modified B e l e m n o s e l l aS , pirulirostrina
as
a
modified
Spirulirostra,Belosepiaas the result of modification of a Spirttlirostrina-l1ke type, etc. I shall present
79) foreshadowsthis change which requires oniy l o d i fci a t i o n s . m i n o r .g r a d u am IV. Finally the centre of buoyancy in sepiids is definitively shifted forwards so that an entirelypas.slve
elsewheremy reasons for considering that such a
maintenance of the swimming position is made
morphologicalseries,which leads from the general
possible.Due to the reductionof the ventral wall, now
type to the specializedtype (cf. Naef 1919,p.20 and
forming the fork, the originally inevitable disturbance
60
d.
e
c l'\
Fig. 38.
Embryonicstagesof recentSepioidea.
a. Median sectionof an embryo of SepiettaowenianaNaef. Note the shell membrane(S) inside the shell sac.Thc latter is divided into two parts: the anterior part (i/s) conespondsto the pro-ostracumand resemblesthat ofteuthoids (Fig. 60). The posteriorpart (11s)would normally containthe phragmocone,which in fact is not formed. A distinct posteriorspine representsthe rostmm. One is temptedhereto imaginethe insefiionof the she1lof a juvenile Spintlirostra. b. More advancedstageofthe samespecies,the posteriorpart ofthe shell sac having disappeared. The muscularmantle reachesthe J(' shellsaconly r ia strandsofconnectivetissue. I nat.size. Do. yolk; Mr. mouth; 1/. inner lip. Ok. upper mandible; Uft. lower mandible; Rl. radular sac; Cg. cerebral ganglion; Pg. pedal ganglion;Sn.bloodsinus;tr/g.visceralganglion; ,St.statocyst; Vc.venacava;Td.inksac; Ma.stomach;Oe.oesophagus; lo.anterior aorta;Ho. Hoyle's organ; Ccr1,2, 3. pafts of the coelom; Sp. posteriorend of body; Mrz. muscularmantle; Go. gonad;Hz.heart; Ms. mantle septum;Ecl. intestine; Mr''. mantle cavity; Vv. venous cross connection;,4f anus; Dr. funnel gland; Ir. funnel tube. Ap posterior aorta; Tk. funnel valve; l/. outer lip; Sr. subradularorgan; Ob. upper buccal ganglion; L'b. lower buccal ganglion; Gg. gangliongastricum;Nl. renal sac;Gr,.genitalvein. c. A moderatelyadvancedembryo in its egg capsule.2{'/,nat. size. 1cf.Cephalopoda,vo1.I. chapter 42). l-5. arms; 4. tentaculararm; 7. pupil; 8. primary lid; 9. olfactory organ; 13. fin; 16. terminal spine; 1B.yolk sac;20. chorion; 21. gelatinousenvelope;22.tip;22. basalparl ofgelatinousenvelope. d. Median sectionof the embryonicshel1of Sepla olJicinali,sL. /. ventral process;2. initial part of siphuncle;Ja. first chamberwith pillars; 30. fourth chamberwith pillars; 4. septalneck of third septum;5. siphuncularhollow of fourth septum;6. p1l\arfor the fifth septum;7. pro-ostracalparl ofconotheca;8. periostracum(dorsalshield). e. Correspondinglateral view. 9. limit of calcified pafi. Ia IVo. septal sutures..f Correspondingdorsal view. 10. angle between lateralridee and ventralDrocess.
6l g.Correspondingventralvicw.IIV.siphuncularpartsofthefoursepta.X.anteriorpoint,offeringasuggcstionofthepro-ostracum. iz.Embryo of Sepia o.f/icirtolisin dorsalview. the shell being visible through the skin. Thrce chambers(1,il, ilI) area\readyformed. T h e d o t t c d p a f t s c o n e s p o n d t o t h e p i l l a r z o n e . 1 , 2 , 3a, r4m . s(tentaculararmsretractcd);5.yolksac;6.eye;T.shellwatl;8-10. sutures,11. Hoyle's organ;-12.teminal spine; 13. fin. d-h 8'1nat. size.
of the (ontogenetically)primary, typical topology (Fig.
Here I will give only a few hints:
62) is no longer required,which restorestotal freedom
Ontogenetic features of the Sepiolidae indeed
(Fig. 35) for the distribution of the internal organs.
suggesta very close relationshipto the Sepiidae(cf.
Flattening of the siphunclealso contributesto this. (The connectionof the soft parls with the gas chambers
Cephalopoda, vol. II, Pl.23 with Figs 15 and 16). It is particularly interestingthat the embryos of this family
is neverthelessconserved).The weight of a bulky posterior rostrum is no longer needed due to the
original shell shapecan be deduced(Fig. 38a);this sac
forward shift of the gas chambers.In this respect
subsequentlydegenerates(Fig. 38a). Before that
show a very well developedshell sac, from which the
Belosepia and Sepia show different gradations;
happens,the site correspondingto the free shell margin
apparentlythe condition achievedtn Sepia is extremely
has shiftedtowardsthe inside of the mantle sac (p. 46);
advantageous.This is illustrated by the extraordinary
Ihe muscular mantle begins to move from this point in
increasein the number of specieswith more or less
typical sepioidfashion(Figs 10, 1l). The primary shell
subtle variants.One gets the impressionhere that a
sac distinctly shows the anterior part correspondingto
goal hasbeenattained.
the free pro-ostracum(Vs) and the posterior part (Hs)
(97) The real courseof evolution from belemnoids
inside which the formation of a phragmoconewould
to sepioids,according to the fossil record, appearsto
seem likely. However, no phragmoconedevelops.It
have taken place at the very end of the Mesozoic
would be interestingto study the closely relatedfamily
period. The results of the potential offered by the
ldiosepiidae(99) for comparison.I supposethat rhis
modifications [of the shell] appearedin essencein the
type, which is more similar to Spirttla, still shows a
Paleocene.The rise ofthe Sepiinaeapparentlyoccurred in the Oligocene.Unfortunatelyit is not possibleto
c l e a r r e m i n i s c e n c eo f t h e p r i m a r y s e p i o i d s h e l l (phragmocone?)during its embryonic development.
trace the geological sequenceof transitional gradesin
Here the anterior part of the shell sac totally
detail, partly due to the scarcity of fossils, which
degeneratesduring postembryonic development, a
certainly does not provide negative evidencefor their occurrencein older or younger strata.See the closing
p h e n o m e n o nk n o w n t o o c c u r i n s o m e s e p i o l i d s (Rondeletiola Naef). As to the anatontical (post-
section.
embryonic) featuresof sepiolids and idiosepiids,it should be emphasizedthat the gonad is only altachedto the stomach'.sincethe genital ligament,which typically
On the relationship of recent Sepioidea to their
(Fig. 62) locatesthe gonadsof decapodsin the area of
fossil representatives.
the siphuncle,is inevitably interruptedby the sepioid penetrationof organs inside the soft body (Figs 9 and
Based on Ihe mvopsid grouping (p. a0) it was hitherto
11 ) , t h e s i t i u a t i o n o b s e r v e d i n t h e t w o g r o u p s
generallyadmittedthat somerecentfan.riliesare closely
mentioned can be explainedby the .former possession
related to the sepiids and spirulids. Since the group
of a sepioid shell. The end result has been conserved
Myopsida has become obsoletedue to the removal of
h e r e , a l t h o u g h t h e c a u s e h a s d i s a p p e a r e d ;t h e
the loliginids and their allies, the questionariseswhere
arrangementof thesesoft parls tells us somethingabout
the remaining families should be placed.Are the shell-
shellform in theirancestors (cf. p.4l).
less families Idiosepiidaeand Sepiolidaereally close
Special elucidation can be expected from the
relatives of the spirulids and sepiids?Comparative
embryonic
anatomical data largely argue in favour of this
D ' O r b i g n y( 1 8 4 5 ,P l . 1 3 ,F i g s 1 3 , 1 4 , 1 8 4 6 ,P a l .U n i v . ,
assumption;see the correspondingchapters in the
Pl. 4) alreadymade use of this feature.But his figures
forthcoming monograph (Cephalopoda,vol. I-Ill).
are much too vagueto make their interpretationuseful.
developmenl of the shell in Sepia.
62 They merely confirm his correct view about the
(p. 111).E. The family Leptoteuthidae (p. 119).F. The
morphologicalsignihcanceof the hump, but he did not
family Geoteuthidae (p. 122). G. The family
recognizethe siphuncularpit (loc. cit., p. 263). A
Belopeltidae(p. 125). H. The family Lioteuthidae(p.
detailed study of carefully removed embryonic shells
132). I. Doubtful forms of Prototeuthoidea(p. 132). K.
of Sepia o.fficinalis (Fig. 38f, g) indeed reveals
T h e M e s o t e u t h o i d e a( p
interestingevidencefor the earlier historT of the type
Trachyteuthidae(p. 136). M. The family Beloteuthidae
accordingto our reconstruction:the anteriorend of the
(p. 141). N. The family PalaeololiginidaeNaef (p.
dorsal shield (Fig. 38x), by its anterior protrusion (x),
141). O. The family Kelaenidae(p. 150). P. The recent
indicatesthe vestigeof a pro-ostracum.A broad (dark)
Metateuthoideaand their relation to the fossil groups
(100) margirral zone is uncalcified. The ventral process
(p. l5a). Q. Review of the evolutionaryhistory of the (p. 161). Teuthoidea
appearsas a specialstructure(1) in the picture,as do
135). L. The family
the lateral edges (10). The lateral view reveals a relatively well marked curvature;on the dorsal side the sLttltresof the early septaare recognizabledue to the
A. Preliminary remarks.
translucencyof the shell. In a median section and also in ventral view, the siphttncular
depressions
Like the Sepia-l1kedibranchiates,the fossil calamary-
(individual septalnecks)are clearly visible; the earliest
like dibranchiatesin recent years have remained
ones are still circular, i.e. not yet dilJerentiatedinto
untouched by systematisfs.The lack of a chambered
fork and hump elements.In the septal neck zone, the pillars are more delicateand more denselyplaced than
shell made them strange-lookingto palaeontologists, while zoologists (confused by many erroneous
elsewhere,so that they appearmore conspicuousin the
indications in the literature, and generally unable to
picture (see Cephalopoda,vol. II, Pl. 20, Fig. 4).
study the fossils themselves)refrained from utilising
Decalcihed sections(unfortunatelyof poor quality) do
fossil forms when studyingrecentspecies,thus missing
not permit a distinction between periostracum and
helpful information. It is our aim here to show how
oslracltm at these stages.I cannot thereforelocate,for
fruitful (102) the combination of zoological and
example,the anterior limit of the ostracum,whether or
palaeontological research can be for the creationof a
not it coincideswith the limit of calcihcation(9).
scientific palaeozoology.
A detailed descriptionwill be given in the context
Fossil teuthoid shells were known already in the
of a comparativedevelopmentalstudy (Cephalopoda,
18'h century (Knorr), but they were probably
vol. II and III). Here only the relation betweenthe
interpreted correctly for the first time by Mtinster
recentand the fossil types can be considered,the fossil
(1828) who comparedthem to the gladius of living
types being elucidatedvia the juvenile stagesof living
squids. The presence of an ink sac indica.tedthe
forms, basedon the lav, of terminal modi/ication (Naef
affiliation with dibranchiates(cf. Buckland 1829).
1917)(cf. above,p. 85-86).
R i i p p e l l ( 1 8 2 9 ) a n d Z i e t e n ( 1 8 3 0 ) r e c o g n i z e dt h e relationship with Loligo. Zieten gave an excellent illustrationof the gladius of a recentLoligo (P1.25,
(t0t)
Figs 8 and 9) for [corrparisonwith] the fossils, and describedthe arrangementof the mantle, gills, ink sac
Part III: The Teuthoidea, or calamary-like
and nuchal region for the sake of further explanation.
coleoids2e.
But early on therewas confusionwith the hook-bearing
N a e f1 9 1 6p,. 1 4 .
Acanthotettthisconocauda (see below, p. 1,78-1,79). Thus, Mrinster(1830,p. aa3) reports:"Sepia.fossilis.
A. Preliminaryremarksand diagnosis(p. Contents: 101). B. On the typical organisationand the developmentof the Teuthoidea(p. 10a). C. The (p. 108).D. ThefamilyPlesioteuthidae Prototeuthoidea
Not only in the Lias of England at Lyme Regis, but also in severalLiassic shalesof southernGermany, fossil ink sacs of Sepia-h.ke forms have been found during the past eight years. I have myself seen some
=
64
near Boll in Wi.irttemberg,some more at Banz and
vol. I, p. 135,and Figs 39c, 61), but their interpretation
Kulmbach in Bavaria,but always amongthe remainsof
is not yet clear, and up to now a systematicstudy of
a Loligo bone belonging to the genus Onychoteuthis
these structures has not been undertaken. (My own
Lichtenst.; in my collection it is therefore labelled
material is still donnant in the form of paraffin blocks
prisca, as alreadynoted in the Appendix Onychoteuthis
awaitingfurther study;but seeFig. 59c1).
fu. aaal on the fossils of Solnhofen (Kefersteins Deutschland von 1828,V/III, 581)".
mean this in the strict senseof our earlier definition (p.
When talking about /oss of the phragmocone,we
Voltz (1835, 1836, 1840)correctly interpretedthe
15). ln fact, in the oldest types we always (in the later
mesoteuthoidsas relatives of the Loliginidae.
ones often) find at the end of the shell a conical to
Quenstedt(1849) considersall the fossil teuthoids, with the only exceptionof Trachytetrthi,s("Sepia")
spoon-shapedstructure, which is always present in
hastiformis Riipp. as calamary-like dibranchiates
It correspondsto the conothecaof a belernnoidand is
("Loliginites");likewisedid Buckland(1836)when he
simply called the "conus". The phragmocone-like
found horny, thin shells (Belopeltis) with "ink sacsof
structuresare to be looked for in the innermostpart of
fossil Loligo" in the Lias of Lyme Regis. They were
its cavity. Strictly speaking,the open conus space is
nearly one foot long and were interpretedas belonging
comparableto the living chamberof a tetrabranchiate;
to an animal which Buckland called "Belemnosepia"or
its predominanceperhaps has a predecessorin the
Sepiotetrthis(cf . 177).
genusCalliconites(q. v.).
advancedembryos of recent representatives(Fig. 61).
Diagnosis: Teuthoidsare decapods(p. 25) in which
On the outsideof this conus, one can often find
Ihephragmoconehas becomerudimentaryand is often
a d d i t i o n a l l a y e r s ( a l s o i n r e c e n t s p e c i e s ! ) ,w h i c h
r e c o g n i z a b l eo n l y d u r i n g j u v e n i l e s t a g e s a s a n
correspondro (105) a sheathwith a rostrLtm.Indeed,
unchambered,spoon-to cone-shapedrelic ("conus") at
such structures are to be expected, because the
the posteriorend of the "gladius", i.e. of the still partly calcified pro-ostracumof fossil forms, - in which the
embryonicrudiment of the conuswould be too delicate to retain its position subsequently.Well-preserved
.fttnnel bay of the ventral mantle margin is limited laterally by distinct, generally prominent angles,- in
fossil remainsof this minor structureare known in the
which the (101) gill a:rls shows a longitttdinal canal
The well-developedpart which is never lacking in a
between the afferent and efferent vessels, which
teuthoid correspondsto the dorsal shield or pro-
communicateswith the mantle cavity via the slits separatingthe gill lamellae, - in which the radular
ostracum. In the younger forms it shows a marked
teeth of the median row always have three points,
clearly resemblesthe typical parts of belemnoids,in
whereasthe accompanyinglateral rows have teeth with
fact so much so (Fig. al) that Voltz, Agassizand others
only two points, one large admedianand a smaller outer pointro,- which always lead a nektonic life
consistently confused them with
("non-stopswimmers").
morphological terms, of course, a gladius (p. 32)
genusP Iesi oteuthis (F ig. 42).
similarity to the recent squids, in the older forms it
belemnoids
(" Belopeltis", "Belemnosepia"). In systematicalways has to be consideredin the light of these older structures;this is the essenceof their interpretation.In
B. On typical organisation and development
particular the "median plate", the "lateral plates",
of Teuthoidea.
median and lateral "asymptotes" or (better) "border lines", "lateral arc zone" and "ventral wall" are to be
As already mentioned(p. 32), a special feature of the
distinguished,to seek homologies with corresponding
teuthoidsis a peculiar modification of the "shell", i.e.
partsof belemnoidshells(Fig. 73).
of its internal parts, which reflects a degenerationof
The free margin of the lateral plate and its growth
some essentialparts. Surprisinglythe loss involves the
lines always (even in the most typical teuthoids)
most peculiar(p. 17) and most ancientelementof the
change direction when they arrive at the (generally
cephalopodshell, namely the chamberedphragmocone.
distinct) lateral asymptote and continue into a zone,
Supposedrelics have been obseled (cf. Cephalopoda,
which in teuthoids is called the "contrs vane". lI
65
Co
Fig. 40.
Gencralmorphologyand topographyof tcuthoidshells.
a. Mantle cavity of Prototeuthis,after removal of the ventralparts of the muscularmantie and slight openingout, in normal position. The funnel apparatusstill showsthe typical character,as do the funnel retractors(Ir), gi1ls(as far as they are visible), arrtts(Afl, intestine(Ed), ink sac (Ib). kidney papillae (Np) and outer gonoducts(G4, the venousappendages (Va) recognizablcthroughthe kidney wall, branchial hearts (Kl), the Musc'ulusrectus abdominis(LIr),the Venaepalliales laterales (Vl),the margin of the posteriorpart ofthe pro-ostracum(G/1),the fin nerves(Fr). CompareFig. 64b and note the consequenccs ofthe degenerationof phragmocone: the the posteriorly advancing muscular mantle and viscera, with a correspondinglengtheningof Ihe Arteriae palliales posteriores (Ap) and Venaepalliales posteriores (Vp), of the fin nerues(Fn) and of the entire posterior abdominal complex;moreovernote the characteristicshapeand position ofthe fins, and the posteriorpartsofthe funnel organ (funnel gland Id) encroachingon the funnel rctractors.Go. olfactory organ;R. funnel tube; tl. funnel pouch; Th. funnel attachmentlVc. vena cava.;Kh. branchialband: Vb. branchialveinl lb. branchial afiery;Pd. pericardialgland; Cl. coelomicpouch accommodatingthe branchialheart:Am. Arteria pallealis ntedianus;,41s. mantle septum;G/2.margin of conus;Rs. Rostrum.The animal is supposed to be a male. 6. Shell of Prototeuthis in situ afler removal of the visccra. Comparethis figure with Fig. 64a for generalconformity and special modifrcationsrelatedto the dcgenerationof the phragmocone.G/. nuchal attachment(gliding surface);Np.Nervuspallialis;St. stellateganglion;x. postcrior limit of dorsal mantle cavity; Kb. insertionof the branchialband; Vl. Venapallealis lateralis: M1t. mcdian plate; ^lp. lateral plate of gladius: Fn. tlnnerve; Vp. Vena palleallis posterior. cntry site in the mantle; Ap.Arterio pallealisposterior; Fl. ftn; Mm. muscularmantle;Co. conus;Rs. rostrum.
66 correspondsto the lateral parts of the conothecaof a
ancestralform, suggestingits superiorefficiency: today
belemnoid (cf. Fig. 72) and smoothly grades into the
severalhundredsof teuthoid speciesoccupy all oceans
"ventral wall" of the "conus", which is homologons
and seasand all zones wrthin them. Most of them are
with the lower margin of the living chamber of an
of moderatesize (gladius length 15-50 cm), but there
Aulacoceras.Theseconditionsare illustratedin Figure
are also giant forms with an overall length of up to 17
a0 (cf. Fig. 64).
m, gladius length over 3 m (Architearlrls).The fossil
Fossil teuthoid shells are often ralher thick (0.5-2 mm) and sometimesheavily calci/ied. But it is difficult
Leptoteuthis gigas attarneda length of more than I tn, as did Tracfu,treuthishasti/brmisl(q. v.).
to determinethe extent of primary mineralisation;the
A typical feature of the earliest teuthoid forrns is
basic matrix is of course conchiolin. In general a
the restrictionofthe very shortfrs to the posteriorend,
layering is easily recognizablewhen revealedby the
where they normally rest on the conus vane (Fig. 42).
splitting of shalescontaining the most well-preserved
The samearrangementis still seenin thejuvenile .fonns
fossils.Accordingto Agassiz(1838)threelayerswere
of most fextant] speciesat the onset of their postembryonicdevelopment. Suchjuvenile stages(Fig.61)
identified by Quenstedt, who described them in "cnttlebones" of the Lias, namely "Loligo bollen,sis"
are also essentialfor the assessment/108) of shell
and "L. aalensis" Zieten (1830, Pl. 25), later named
morphology. They show the contrs as a much more
"Loligosepia" (1839,p. 163). (The (106)original
well developedstructurethan in the adults and reveal
specimensare housed in Tilbingen, where I have
its typical relationship with the muscular mantle (p.
examinedthem). fQuenstedt]distinguishes:L a thick,
22), which surroundsits aperture(Fig. 61a). It has a
white layer of mother of pearl; 2. a brown, still horny
rounded shape,and a virtually inhibited transition to a
dorsal layer, and (107) 3. a covering that is not very
protoconchof a belemnoid could be imagined.(For
sharply demarcatedfrom 2. Morphologically 1in the
further illustrations see, in addition to Fig. 39, the
senseof p. 13) the three layers can be interpretedas
monographCephalopoda, vol. I, p. 135-162).
hypostracum, ostracum, and
periostracum,
respectively.I have found ample confirmation that the older, more robust gladii show all the threemain layers
C. The PrototeuthoideaNaef 1921
of a cephalopodshell. (Quenstedt'sideas about the
(System.p. 534).
generalorganisationof the shell architecturewere not so clear3r).He did identify the "parabolic fields" with
Diagnosis: Prototeuthoidsare fossil teuthoids(Lias to
the "bow region" of the belemnite shell; he calls the conus vane "wing fields"::. - His theory of
Cretaceous) in which the median plate (which in
stratificationoften served,in an ill-conceived fashion,
or less heavily calcified gladius has a very blunt
to assume here a transition to the humo of the
anterior end (occupyingat least one half ofthe mantle
cuttlefish.
width), without a ventrally concave,broad median
The typical organisation of the so./i body of
generalis sharplydelimitedby asymptotes)of the more
keel. Insteadvery fine lines, groovesor coarserribs
teuthoids is illustrated in Fig. 39. Well-preserved
may occur, and the shapeof the longitudinal lines and
fossils(e.g.Figs 42 and 43) prove that the generalbody
growth lines always indicatesa well developedconus,
shapeof the fossil fonns must indeedhave been squid-
which is usually broken off.
like. Comparedwith the belemnoids,the degeneration of the phragmocone(as in sepioids)is conditionedby an increaseof the muscular mantle.The resultof this
These forms are closely related to the type describedon p. 103 [i.e. Fig. 39]. Their essentially correct interpretationdatesback to Quenstedt(1839,
metamorphosiscan also be viewed as a shift of the
1849)who suggestedthe homologiesof their structures
morphologic-ecologicalequilibrium: along with the
(1849, p. 503) with those of belemnites,assuminga
gas-containing phragurocone a
pointed conus and distinguishinga "bow region", a
compensatory
weighting by a rostrum is abandoned (see under
"parabolarfield" or "middle held" (our median plate),
Belemnoidea);the animals have to rely on Iheir active
"fields of hyperbolic lines" (our lateral plates) and
energy rather than on a passive, cumbersome
"wings" (our conusvane).
apparatus.The new type has completely replaced the
A tvoical feature of these shells is a marked
Fig. 41.
Prototeuthoidshellsfrom the Lias e flower Toarcian]of Swabia.r/2nat. srze.
a. Paraplesioteuthishastata,drawn (with somereconstn.lction) liom a specimenin the BavarianStatecollcctionsin Munich. (This is the original specimenfrom Holzmadcn,describedby Miinster, Bcitr. 6. P1. 14, Fig. 4). Note the secondarynarrowing comparedto D and the densestriationon the marginalpartsof the medianplate at -r. Dorsal vien . b. Poraplesioteuthissagittato,carcfully reconstructedfrom the original specimendescribedby Mtinster (Beitr. 6, Pl. 7, Fig. 3) in the Bavarian StateCollections(public collcctions).Posteriorand anterior ends reconstructedon the basisof the growth lines visible on the liagment.Dorsal view with (ruptured)ink sac.Collectednear Boll. c'.Paraplesioteuthissagittato var. mogno, drawn from a fine specimenin the collection of B. Hauff (Holzrnaden).Similar to thc previous form, but somewhatmore slenderand much larger. To be reconstructedlike the previous specimen.The broken lateral marginsarereconstmctedon the basisof the grou'th lines of thc survivingpart. d. Lioteuthis problematica, drawn from a shell in the Univcrsity of Tiibingen collections. labelled no. 6146 (cf. p. 132). This specimenrcpresentsa peculiartype *'hich showsno closerelationshipto any known genus.Note thc bandslying betweenthe lines_i, and a; the natureof thesebandsis llot quite clear.On the basisof the growth lines they can eitherbe interprctcdas a marginalpart of the medianplate (like,t in Fig. a), or as atypicallateralplates. 1. paramedialribs:2. median groove,partly with a very delicatcridge; J. smear-like.irregulardepositson the dorsal side; 4. reinforcedpart of lateral plate; 5. median asymptote;6. median part of median plate with (in this species)particularlydensearcuate stripes;r. ink sac;-r,.probably the indistinct mcdian asymptote;r,. longitudinal ridge; z. central rib; a. lateral asymptote;c. conus vane.Dottedlines mark reconstructed Darts.
68 similarity to the pro-ostracaof belemnites,a similarity
associatedwith the free margin of the shell, as would
that misled voltz (1835, 1836, 1840)to identify them
be expectedin a prirnitive state(p. 108). In the adult
with belemnites.To confirm his (otherwisecorrect)
the musculatureof the mantle on either side continues
idea of belemnite shell architecture,he neededfossil
tunderthe lateralplates(Figs 42 and 43) and apparently
remainsof the pro-ostracum,and sincetherewere none
always extendsto the median plate. Here we see the
available to him, he found them in the shells of
oppositeol what we find (Fig. 7c-e) in recentteuthoids
prototeuthoidsthen known under the name of (109)
(p. 158);this could be relatedto the limited flexibility
"Onychoteuthisprisca" Miinst.33.This neverthelesshad
of these [i.e. fossil] shells. Perhapsthe shell parts involved are not strictly homologouswith one another icf. Trachvteuthis\.
the positive effect of an early recognition of the homologiesbetweenteuthoid and beiemnoidshells.On the other hand. Voltz obscured some facts about belemnite shells: to impose his view, he arbitrarily reduced the observed growth lines3aof Belopeltis
D. The family PlesioteuthidaeNaef 1921
aalensis (Fig. a7) to make them fit in with the
(Systemp. 534).
belemnitetype (1840, Pl. 4) (d'Orbtgny's Belemnites aalensisarosein the sameway {1842,Pal. Fr. Jur.,Pl. 3 - 4 )) . A p p a r e n t l y V o l t z a n d A g a s s i z m u t u a l l y
Diagnosis: More or less slenderprototeuthoidswith a conus vane that is restrictedto the posteriorend and is
e n d o r s e de a c h o t h e r i n t h i s v i e w . A g a s s i z ( 1 8 3 5 )
bent down to form a pointed cone, grading anteriorly
published an (erroneous)note on a belemnite with
into the relatively broad, gradually tapering lateral
distinctremainsof the pro-ostracum(cf. p. 168).(This
plates,
in fact was a broken phragmocone with an ink sac
in which the rnedian plate bears a simple or
inside and with the rostrum of a paxillose belemnite).
double median rib and narrow lateral bands clearly demarcatedfrom the broad central part. - The hns are
Later on (translatedby Buckland, 1838, explanationof
very short, subterminal, resting on the conus. -
Pl. 28) he stated:
Nominal type: Plesioteuthis prisca (Fig. a2).
"The differencebetween these gladii, which are
The unity of this new group is easily recognizable
madeup of three layers,and the gladii of living Loligo
from a comparisonof Figures 41b and 42, even if
species, which have only one layerl5, and their
intermediate forms (Fig. 4la) are not taken into
correspondencewith the continuation of the shaft of
c o n s i d e r a t i o n .P o s s i b l y t h e s h e l l s o f t h e g e n u s
the belemniteillustratedin PL 44', Fig. 7, convinced
Paraplesioteuthis from the Lias belong to the
me that the so-calledLoligo
immediateancestorsof the Upper Jurassicfonns. But a
aalensis is
the
prolongation of the belemnite, as is clearly
step-wiseprogressis out of the question.
recognizable in this specimen. This circumstance causedme to changethe name Belemnites(which was
The genusParaplesioteulrisNaef 1921
previously used for the well-known frostrum] of this animal) Io Belemnosepia, thus indicating the close
(Systemp. 534,539).
relationshipof this animal to the cuttlefish...".fNote:
Diagnosis'.More or lessrobustplesioteuthids(from the
the name Belemnoseplahas been suppressed by the
rczNl.
Lias), in which the median rib of the shell is double anteriorly and the central part of the median plate
For a solutionof this confusion seethe sectionon Belemnoidea. Agassiz notwithstanding, Buckland
dorsally carries severalsystemsof parallel, retrograde transversestriations.- Principalspecies:
(1835) and Quenstedt(1839) clearly distinguishedthe gladii of prototeuthoidsfrom the supposedbelemnite pro-ostraca.(Il0)
The same is true of d'Orbigny
(1842, 1845).For us it is essentialfto recognise]the strongequivalencebetweenthe pro-ostracumand (111) the prototeuthoid gladius. In several genera I found peculiar features in the insertion of the muscular mantle: Only in the region of the conus does it remaln
012) 1. Par aplesiot euth is sagittuta (Miin ster 1843). Herebelongs:Geoteuthis sogittatdMiinst.(Beitr.6, 1843,p. 72, Pl. 7, Fig. 3), aswell asthevarietydescribed there(P1.8, Fig.4). Belemnosepia sagittatad'Orb.(1845and 1855,p. 4 3 8 . 1 8 4 5P a 1U. n i v .P l . 2 7 . P a l .E t r .P l .2 4 1 8 5 0P r o d r . 1,p.242). Loliginitessogittatus (18a9,p. 516,Pl. Quenst.
=
69 Fig. 42. - Plesioteuthisprisc'a, drawn from different spccimens from the lithographic limestones of r/2 Bavaria. nat. size. a. Gladius with the impressionof the whole animal,
'", nl t",Yf LT l
,Ji
!a
.:i,i ' i/ \,1 i . ' . . . i i r , ' , ' i :: ;
i
'.
e.
i
;.r"::'":
drawn from the original specimen from Eichstiitt ,;:i.
describcdby Zittel ({Broili}, Grundziige1915.p. 583). Apart from minor additions on thc flanks of
t
the pafts drawn are those safely recognizableon the
-Va
i
.
l
i
I
,
,
,
the mantle,no reconstrxctionhas beenmade. so that
:
'
1 . i.
specimen.Note the very distinct fins, the terminal :UX.
:
apex ofthe conus,the ink sac and duct, the cephalic
KK
r,. fiLl
cartilage and eyes, the upper mandible and in particular the arms, the dorsal (Do), dorsolateral (Dl) and ventrolateral(fD of the left side being identifiable.The arms bent laterally (Lt and Rt) are
o.
not very distinct but can be assumedto representthe tentacles.and the arm bent backwards (Va) is plobably a ventral arm. Overall one can recognizea
Y*-,;'
t1,pica1dibranchiateshowing the characteristicsof
n-.,--,t9+-,; !
the youngestteuthoids(Fig. 6 I ), in dorsalview.
{;=i1'}
b. Completc shell in ventral view, drawn from a
t1t
Sr Sr srr l.r ,f
N"l"{s, I \.'l /
beautilul specimenin the BavarianStateCollections (like ri).
: l'!i r )
Mr. middle field of the median plate,
l i
darker in colour at z: S/. lateral field of median plate; Sr. dividing longitudinal rib, terminatingat z;
i
r c.'
5p. lateral plate; Z.s. fine longitudinal lines; ,t. incurved oblique striations(marked as dotted lines. since mostly indistinct); 1. anterior end of central
,"l
rib (Mr):7'b. ink sac; c'f conus vane; Ia. mcdian asymptote;Aa. lateral asymptote;Co. ventral wa11 of conus,reconstructed;Ro. rostmm. Compare the incompletefigurein Zittel (1885.p 519).
z
c'.Lateral view ofthe posteriorend ofthe conusofa
lb
specimenfrom the Hiiberlein collections (Munich),
d
posterior dotted part added.Mm. muscular mantle; note its insertion on the conus margin, whereasthe rnuscle bands cross the edge of the pro-ostracum. Solnhofen. r/. Young individual of Plesioteuthi.r from Eichstiitt; specimenin the Munich collections. e. Posterior end of a, reconstructedfrom several specimens.Note the insertion(Fg) of the fins on the
11a
conus and the overlappingof the lateral plates with a t
the muscular mantle, which is typical of
Jcr
prototeuthoids tp. I I I t. /. Isolated (upper) mandible tiom the lithographic stoneofNusplingen.cf. Quenstedt1858(Jurassic),
i.
P1. 99, Fig. 22. The specimen is in the Tiibingen collectionsand is labelled "Saepia beak"; it very likely belongsto this species.It is a typical decapodmandible (cf. Cephalopoda, vol. 1, Pl. 17) and proves at least one thing, namely that theseforms, which are so similar to recent forms, already occurredin the .Iurassic(cf. p. 25). I havc seensimilar examplesassociated with an animal,but noneof them was so well preserved.
70 3 5 ,F i g .3 1 8 5 8p, . 2 4 4 .1 8 8 5p, . 5 0 7 ) . I b i d .C h e n u1 8 5 9p, . 4 1 ,F i g .1 1 7 .
below p. 161).I havenot, however,beenableto seeits
This very characteristicshell form is particularly
This speciesseemsclosely relatedto the following
useful as a basis for a general rnorphology of the teuthoidgladius(cf. Cephalopoda, vol. I, p. 137).But
ventralmargin.Note the ink sacl one from the Lias e fl-ower Toarcian], but apparently at a slightly higherhorizon,henceyounger.
at the presenttime it is not the oldestone. According to information from Mr
B. Hauff of Holzrnaden,
Belopeltisaalensis and Geoteuthissimpler occur there in the Lias e fl-ower Toarcian],in earlierrocks than the
2. Puraplesioteuthis hastata (Mrinster I 843). Herebelong:Geotetrthis hastataMiinst.1843(Beitr.6, Pl. 8,
number of well preservedspecimensis rather small.
Fig. 3; P1.14,Fig. 4, p. 13).Belemnosepia hastatad'Orb. ( p . 4 3 9 ) ; ( P a 1 . 2 8 , F i g .l , P a l .E t r . 1 8 4 51, 8 5 5 1846 U n i v .P l . Pl. 25, Fig. 1). Belopeltishastata"ibid. 1850(Prodr.l, p.
They vary in their proportions,some are more slender,
242).
present species.The latter is not abundantand the
others are more robust, or more delicate; I would
Comparedwith the previous species,this form is
distinguish different species if there were not
strikingly delicate and narrow, similar in outline to
intermediateforms between the extremes,and sexual
Plesiotetrthis,with particularly densearcuatelines on
dimorphism also has to be taken into consideration.
the lateral fields of the median plate. Otherwise,in all
The main forms are illustratedin Fig.41b and c; the
essentialfeaturesit is very similar to the previous
variety magna probably measuresmore than 30 cm
species.The changein form of the anterior end results
(afterreconstruction).
in nearly parallel lateral margins, which can be
The median plate is acutely triangular, the acute anglepointing to the rear.The most posteriorpart bears
interpretedontogeneticallyas a lengtheningwithout a correspondingbroadening.
a strong dorsal, median rib (reminiscent of Plesioteuthis)which splits into a pair of paramedial
The genusPlesioteuthisWagner 186016.
ridges. They progressivelydiverge anteriorly, flanking a narrow groove. Finally the whole structure flattens
Wagner (1860, p. 36) first distinguishedthis best
out. On either side of this complex median structure
known fossil teuthoid as a separategenus, but it had
lies a field with a transverseto obliquely rippled
alreadybeen recognizedas an apparentlyseparatetype
surface.The strips run medially and posteriorlyand
by Voltz (1836) and by Mtinster (1839). fNote: The
form severalsystems:a filore oblique one anteriorly, a
genuswas actuallyfirst usedby Wagnerin 18591.
less oblique one posteriorly; in the central area these
Here belongsonly one species:
systemsintersect one another. In the anterior part, a dorsal view additionallyrevealssmear-likeshadows,
Plesioteuthisprisca (Rtippell I 829).
which appearlike a bluish-greyish-whiteenamelcover.
First describedas "Loligo" prisctts Rripp. 1829,p. 8,
The lateral parts of the median plate are delimited by
Pl.3, Fig. 1. (Securelyidentifiableillustrationof the
sharplines and show a conspicuous,regular pattern of arcuate strips that join the lateral asymptote. These
conus vane, median rib, longitudinal ridge of the
structuresare growth lines that can also be seenin the
muscularmantle and fin impressionare presentin their
medianfield of the most anteriorpart of the shell, so
typical arrangement).Subsequentlyvarious names
that a reconstructionof the missing anterior margin is
have been introducedinto the literature,mostly based
possible.
on incompletelypreservedspecimenswhich looked
anterior part of the gladius. Ink sac, parts of the
The lateral plates taper anteriorly and are indeed
more or less unusual,but sometimesbasedon quite
very narrow in the middle part of the gladius (in
(Il5) typrcal specimens.Thus, Mtinster (1830, Jahrb.
generalthe delicatemarginal portion is (113) partly
p . 4 0 4 , 4 5 8 ) a l r e a d y r e p o r t e do n O n . v c h o t e u t h i s
destroyed).Posteriorly they gradually grade into the
ongLtsta,a name subsequentlyadoptedby Voltz (1835).
conus vane and, like the latter, show a dual pattern of
Later Mtinster ( 1846)3?described them
stripes:lines parallel to the free edge are growth lines,
Acanthoteuthis angusta (Pl. 4, Fig. 1-3) along with
while othersconvergein an acuteangle pointing to the
other noveltiesand figures.Unnamed:Pl. 4, Fig. 4-6,
conus,so that the latter can be easily reconstructed(cf.
P I . 5 , F i g . l - 5 , P I . 6 . ,F i g . 3 . A c . l a t a ' P . I.6,Fig.5.lc.
as
71
l . A c . s t r b o v a t aFi i g . 2 . A c . s u b c o n i c a :F t g . 3 . A c .
Q u e n s t e d(t1 8 a 5 ,p . 5 1 8 , P l . 3 5 , F i g . 3 - 4 ) r e c o g n i z e d only "Loligo" priscus and apparently called the
acuta: Fig. 4-5 (uv.). The confusedsynonymyof this
validity ofthe otherspeciesin doubt.(Seealso 1885,p.
speciescan be taken from d'Orbigny (1845, 1855);he
508, Pl. 39, Fig. 7-8; herethe conusis drawn as being
listed it underthe followine 5 names.with indications
ventrally open;he did not recognizethe homology with
of fufiher synonymies:
the conusof belemnites).
tricarinata'.Pl. 6, Fig. 6-7; Ac. semistriata:Pl. 7, Fig.
l . A c a n t h o t e u t h i sp r i s c a 1 8 4 5( 1 8 5 5 ) ,p . 4 0 9 , P l . 2 8 . Here belong the bodies and arm crowns of our AcanthoteuthisspeciosaMtinst. (Fig. 1-3 {p.180}, plus a gladiuswithout the conusof Ples. prisca (Fig. 4). Both forms also occur as synonyms,of which I here cite only those of Plesioteuthis:Loligo prisctts Riipp. 1829, OnychotettthisangustaMiinst. 1830, KelaenosagittataMtinst. 1836MS., Onychoteuthis sagittata Miinst. 1837 (Jahrb. P. 252), O. angusta
Miinster thought his supposed species were "teuthoids" with hooks and therefore named them "Celaeno",laterhe followedWagner(1839)and called them "Acanthoteuthis".(Seeunder Belemnoidea).The stomachsand coproliteswhich apparentlybelong to the specimensindeedcontainhooks along with the remains of shells.Also somehooks have apparentlybeen found at the anterior ends of the "sacs" (Beitr. l, p. 104, VII,
ibid., O. tricarinata ibid., Acanthoteuthis brevis
p. 57). Wagner ( 1860) already observed that
Miinst. 1842 (Beitr. 5, Pl. 1, Fig. 3), Ac. pt'isca
Plesiotetrthisnever bears (117) hooks that could be
d ' O r b . 1 8 4 6( P a l .U n i v . , P l . l 9 - 2 0 , P a l . E t r . , P l . l 6 -
consideredas the annour of their arms; figure 713, p.
t7).
517 in Zittei (1885) is clearly misleading:the hook-
2 . O m m a s t r e p h ecso c h l e a n s1 8 4 5( 1 8 5 5 ) ,p . 4 1 7 . P l u s Orrychoteuthis cochlearis Miinst. 1837, p. 252. -
shaped lines are either incomplete renderings of the
Ommastrephes cochlearisd'Orb. 1841,C6ph.Acdt.
that may be observedin such impressions,as we have
l n t r o d .p . X L , i b i d . 1 8 4 6 ;P a l .U n i v . ,P l . 2 4 , F t g . 2 , p .
seen (p. 122) tn other teuthoids. I interpret them as
2 0 1 ; P a Ld t r . ,P l . 2 1 , F r g . 2 .
t r a c e s o f m u s c u l a r s t r u c t u r e s ,e . g . s u p p o r t s o f
3.Ommastrephea s n g t t s t u s( p . 4 1 5 , P l . 3 0 ) . P l u s On.,-chotetrthisangusta Miinst. 1830, Jahrb. p. 404 and 1836,p. 250, 630. (lI6) O. lichtensteinii1.831, Mtinst. MS. O sagittata Miinst. 1837, p. 252 (not Lam. 1799 ). O. angusla Miinst. 1837,p. 252. Ommastrephesangustltsd'Orb. 1846 (Pal. Univ., Pl. 2 3 , F i g .9 - 1 1 ,p . 4 l 2 , P a l .E t r . .P l . 2 0 .F i g .9 - 1 1 ) a , nd 1850(Prodr.1.p.347\. (p. 416). Here belongs 4.Ommastrephesintermedlars Onychotettthis intermedla Mrinst. 1837, p. 252. Ommastrephesinterntedlas d'Orb. 1841 (C6ph. Ac6t. lntrod.p.XL) and 1846(Pal.Univ. PI.24, Fig.
arm impressions,or they representtransversegrooves
protective membranesas observed in all decapods (Cephalopoda, vol. I, p. I 18). Newer textbooksand handbooksoften show figures of Plesioteuthis,(cf. e.g.Keferstein1866,Pl. 130,Fig. 7-8, Zittel, 1885,p. 519),but noneof theseillustrations provides more than a vague representationof the overall morphology. I therefore draw attention to Figure 42 which offers the possibility of a more rigorous comparisonof teuthoid shells with belemnoid shells. Plesioteuthisprisca is probably the most abundant dibranchiatein the lithographic limestones(i.e. Malm
l , P a l .E t r . ,P I . 2 1 ,F i g . 1 . Loligo
[E/] e) of Bavaria fl-ower Tithonian] and Wiirltemberg
sttbsagittata Miinst. 1836 (Jahrb. p. 582)38,ibid.
[Upper Kimmeridgian]tu. I have not seen further
1 8 3 9( p . 3 7 5 ) ,i b i d . 1 8 a 3f u . 1 0 7 ,P l . 1 0 , F i g . 3 ) .
specimenslike the one in Figure 42a; but there are
(This form shows a broad vane similar to the living
many moderatelywell presenredexampleswhich allow
genus Enoploteuthis; in reality it is only the
one to intelpret poorly preservedspecimens.From a
compressedmantle sac). See also d'Orb. 1845, p.
comparisonof all of them the animal can be described
3 9 8 ,P l . 1 9 ,a n d 1 8 4 6( P a l .E t r . .P l . 1 5 .P a l .U n i v . ,P l .
almost as completelyas most extant relativeshad been
18), original specimenin Munich. The "species" 1
describeduntil recently.
5.Enoploteuthissubsagittata (p.
358)
:
and 3-5 are still recognizedby Fischer1887,p. 348. Seealsov. Biilow 1920.o.263-266.
The
shell
clearly
resembles that
of
Paraplesiotetrthissagittata (Fig. alb) in its overall
72 aspect,but is more slender.The similarity concefirsthe
Plesiotettthisprisca; we have not much precise
subdivisionof the medianplate into median and lateral
information and therefore have doubts about the
fields, the narrow lateralplates,the shapeand sculpture
usefulnessofa separategenus.
of the conus. Posteriorlythe conus bears a distinct, small rostrum (which is probably also present in
Dorateuthis syriaca Woodward I 883.
Paraplesiotenthi.s sagittata); it is sometimesembedded
The original illustration provided by the author (Pl. I,
in a way which allows its original form to be
Fig. 1) showsa small, slenderdecapodrneasuring7 cm
recognized.The three longitudinalribs are conspicuous
in length; it was found in the Upper Cretaceousof
in addition to finer longitudinal lines. The central rib
Sahel-Alma fin the Lebanon] and could be identical
occupies more than the posterior two fifths and is a
with the speciesof our Figure 50, which is from the
rounded rod tapering at both ends. It therefore must
same site. The general form is that of Plesioteuthis
have been fonned by secondaryadditions. The lateral
prisca (Fig. 42); but as far as can be seen, a strong
ribs, which are most distinct in the anterior third,
central rib extendsto the anterior end of the gladius,
separatingthe median fleld from the lateral field, are
and the arms appearstiff and pointed, as in Figure 50.
much weaker. They also taper anteriorly and
Apparently there are traces of a (I19) tentaculararm.
posteriorly.The (118) lateral fields lack the sharp
[Note: The tentaclesshown in Woodward's plate were
arcuatepattern observedin the previous genus,but
an imaginary additionl. Securely observableare eight
otherwise they have the same shape. In particularly
arms with hints of suckers, the head with eyes and
w e l l p r e s e r v e d s p e c i m e n s ,a d e l i c a t e r e t r o g r a d e
mandibles,the mantle sacwith gladiusand ink sac.The
transverse striation can be observed in the median
broadeningof the posteriorend resemblesthat of many
fields (x).
specimensof Plesioteuthis and is due to the fins and
The correlation with the soft parts is most
conus. The shape of the latter is not clearly
interesting,as shown in Figure 42c and e. The inserlion
r e c o g n i z a b l e ,h o w e v e r . T h e c e n t r a l r i b a p p e a r s
o f t h e m u s c u l a rm a n t l e ( p . l l l )
and of the fins is
reinforced. Woodward assumes broadened lateral
clearly recognizable, which is irnportant for the
plates in his reconstruction,so that Dorateulftls would
confirmation and completion of our reconstruction deducedfrom the morphology of recent teuthoids (p.
be closer to Leptoteutftis. But the lateral plates are apparentlyindistinct in the fossil itself. - ln its details,
22 and 34). However, I have never seen the fins as
the tentative reconstruction is arbitrary or shows
sharylyoutlinedas in Fig. 42a; the mantle sac also is
The accidentaloutlineswithout a critical assessment.
generallymore compressed.
following forms appearagaincloseto Plesiotettthis:
The outline ofthe head is that of a slenderteuthoid in general.Figure 42a very distinctly showsthe eyes,
The genusStyloteuthisFritsch 1910.
the head cartilage (or optic ganglia) and the upper mandible. The arms are short, stout and unequal in length,which increasesfrom dorsalto ventral. They do not show distinct suckers but some imprints left by suckersor cirri; they could be mistaken for hooks. Whether the irregular impressionslabelled Lt and Rt
The original description and illustration of the respectivespecieswas not available to me. Btilow (1920) describedthe following features:longitudinal axis with strong,roundedkeel, posteriorlypointed.The following speciesare cited: St.catrdateFritsch,p. 13,Pl. 5, Fig. 2
representtentaclescannotbe said with certainty(cf. the arm crown in Fig. 50). - The gladii of this speciesin general measure 20-25 cm in length, they are rarely
St.convexaFritsch,p. 12,Pl. 5, Fig. 1.
slightly longer or shorter.This probably correspondsto
Both are from the Upper Cretaceousof Bohernia. "Acanthoteuthis " maestrichtensis Binkhorst 1861
the normal size of the adult animal, which varres
(p. ll,
widely in dibranchiatecephalopods.(cf. Cephalopoda,
Maestrichtprobablyalso belongsto Plesioteuthis(cf.
vol. I, chapter7). The following speciesis close to
Z i t t e l 1 8 8 5 p, . 5 1 9 ) .
P 1 . 5 d , F i g . 4 ) f r o m t h e C r e a t a c e o u so f
aa / J
E. The family Leptoteuthidae Naef 1921 (Systemp.534). Herebelongs: The genusLeptoteuthisH. v. Meyer 1834. Only one species: Leptoteuthis gigas H. v. MeyeCo. Synonyms:Leptoteuthisgigas H. v. Meyer 1834,p. 292, Ibld. 1836, p. 56, Acanthoteuthisorbignyana Miinster 1846, Pl. 7 , Frg. 6; Acanthoteuthislata Miinster 1846, Pl. 6, Fig. 4: Acanthoteuthis gigantea Mtinster 1846 (VII). Pl. 8. Unnamedform of shells: Mrinster1846(Vll), Pl. 6, Fig. 1-2; (120)Leptoteuthis g l g a s d ' O r b . 1 8 4 6( P a l .E t r . P l . 1 5 . p . 3 6 3 ^ P a l . U n i v . Pl. 12); Loliginites alattrs Fraas 1855, p. 88; Leptoteuthisgigas (121) Wagner 1860,p. 19-26,P|. 2; Leptoteuthis gigas Chenu 1859,p. 31, Pl. 99 (unrecognizable),Leptoteuthisgigas Keferstein 1866, p . 1 4 4 3P , l.130F , i g .6 . This species is probably as abundant as Plesioteuthis,but given its size (the shellmeasuresup to about I m in length) destruction of many parts is generallygreatersinceburial must have been slower.A small specimen figured by Crick (1915, as "Plesioteuthisprisca") is therefore of special interest. Its generalform clearly resemblesthat of the previous species;but the animal is not as slender,the fins are drawn aparl more markedly,almostband-like,tentacles are recognizablewith certainty. The gladius shows a similar subdivision of
the median plate as
Plesioteuthis; this originally (prior to the general revision of fossil dibranchiates)led me to place both generain the same family (Cephalopoda,vol. I, p. 47
Fig. 43.
and 143). A
reconstructed from a photo ofa Solnhofcnslab (Crick 1915, describedas"Plesioteuthispriscct").I,a nat. size.Ofspecial
more detailed inspection reveals
considerabledifferences:the lateral plates are broad, leaf-like, the conus is blunter. The lateral fields of the median plate are clearly narrower than the median
Young individual of Leptoteuthl,sglgcrs.
interestarc the fu1lypreseruedimpressionsof the fins (fl and tentacles(I) which arc missing from most specimcns.The slab showsthc animal as a perfect image so that the arms,the
fields, and instead of a rod-like central rib there is a
head,the mantle sac,the fins. the conus(with a rostrum?)can
band-like, flat rib with anteriorly convex transverse
be safelyreprcsentedin the figure. The eyesand suckershave
striae,which may appearas a sharpridge only close to
beenadded(cf'.Cephalopoda,vol. I. p. 143 and 148).
the posteriorend. The strengtheningridgesbetweenthe median and lateral plates are less distinct, rather flat, well developedonly in the middle and posteriorparts.
In the relatively blunt conus of iarge specimens(as in
Anteriorly convex transversestriae are presentin the
Fig. a4) transversefolds, which might suggestthe
rnedianplatebut arernorewavy in aspect.
presenceofchambers (cf. p. 104) are probablyaftefacts
74 completely compressedand spread out in a single plane. The gladius is sufficiently well preservedto permit secureidentification (like in Ftg. 44, but with a less markedly blunt conus). The muscularmantle is recognizablefrom the transversestriation and the rough outline. What had been interpretedas the funnel is merely a torn piece of the rnantlesacwhich has been displacedlaterally (the animal lies ventral side down). The overall shapeof the headwith remainsof the (122) mandiblesand four arm basesis also recognizable. Hooks have been reportedon the ann bases.There are indeed transversestructuresthat could heve been causedby suckers,their basal cushionsand protective membranes. But what today seems so strikingly distincton this specimenis a resultof preparation. The blue-grey remains of the animal have been enhanced during preparation by exposing the surrounding calcareousmaterial which is of lighter colour. Of courseany imaginablepattern can be producedin this way, and we can therefore no longer determine what was really visible prior to maltreatment. Hooks sirnilar t o t h o s eo f A c a n t h o t e u t h i s ( F i g . 9 l ) a r e o u t o f t h e question. Only soft parts have caused the dark markings reminiscentof such hooks; this has to be ernphasized,to avoid a new misleading assumption about a "Kelaeno with hooks" (cf. Fig. 56, and vol. I, p. A7). Cephalopoda,
F. The family GeoteuthidaeNaef 1921 (Systemp. 534). The genusGeoteathisMrinster 1843,pars.
Fig. 44. LeptoteLtthi.\glgas Meyer 1834, drawn from the original specimenfrom Daiting (now in the Munich research collections, Hiiberlein collection) described by Wagner (1860,Pl. 21. I12nat.size.Left side reconstructed. ll. inner
The speciesbelonging here, from the Jurassicand C r e t a c e o u s .h a v e b e e n g i v e n v a r i o u s g e n e r i c d e s i g n a t i o n s :" B e l e m n o s e p i a " A g a s s i z 1 8 3 5 , d'Orbigny 1845 (p. 433), Loligosepla Quenstedt1843,
asymptote;Ae. otler asymptote.Note the curved lines
Belopeltis Voltz 1840, PalaeosepiaTheodori 1844,
coveringthe growth lines of the medianplate.
Loliginites Quenstedt1849. They were confusedwith belemnite soft parts, belopeltidsand other decapod
causedby compressionduring burial. - During the Late
remains.We shouldconsiderGeotettthissimplex(Yoltz 1840 : Geoteuthis/afa Mrinst. 1843 : G. orbignvana
Jurassic these animals may have played a role
ibid.) as the leading species;it was lirst described
comparableto large extant forms llke Loligo vulgaris,
underthis narne.(Firstillustratedin Buckland1838,Pl.
which are the same size. In Tribingen (University
30).
collections)I have been able to study a large,very well p r e s e r v e d s p e c i m e n .I t s h o w s t h e w h o l e a n i m a l
Diagnosis'. Bulky Prototeuthoidea.in s'hich the broad conus vane extendsfar forwards. thus flankins
=
75 the posterior half of the pro-ostracum (as a leaf-like s t r u c t u r e-) .L i a s - M a l m . I have only recently realized the profound differenceof this shell form comparedwith Belopeltis aalensis(p. 125).ln Cephalopoda (vol. I, p. 47) these types are still united. (I separated them in loc. cit., p. 143, but this was done using untenablenameswhich reflect a confusednomenclaturalhistory). The essential f'eatureis the structureof the conusvane, which places theseforms (123) betweenParaplesioteuthis(Fig. 41b, c) and Belopelrls(Fig. 51 l54l). Geoteuthissimplex (Voltz 1840). Voltz interpretedthis type of shell as the pro-ostracum of an unknown belemniteand named it "Belopeltis" simplex(1840,Pl. 2,Fig. l-2; herewe alsoplacehis B. e m a r g i n a t a ) . I n 1 8 4 1 ( J a h r b .f . M i n . , p . 6 2 5 ) t h i s interpretationwas further consolidated.Miinster ( 1843, Beitr. 6, Pl. 7, Fig. 1) calledthe sameform Geoteuthis lata and (Pl.l , Fig. 2') G. orbignyanaal;d'Orbigny ( 1 8 4 5 , 1 8 5 5 )l i s t e db o t h " s p e c i e s "a s B e l e m n o s e p i a (p.436,438)and illustratedthe first one (P1.31, Fig. 1). I n 1 8 4 6( P a l .U n i v . ,P l . 2 5 , F i g . 1 , P I . 2 6 , F i g s I a n d 3 , a n d P a l . E t r . ,P l . 2 2 , F i g . 1 , P I . 2 3 , F i g . I a n d 3 ) h e illustrated both of them.
Quenstedtrecognisesonly " L o l i g i n i t e ss" i m p l e x( 1 8 4 9 :P l . 3 3 , F i g . 6 - 7 ,p . 5 l l ; P l . 3 4 , F i g . l ; 1 8 8 5 ,1 8 5 8 :P l . 3 9 , F i g . l 0 ) . C h e n u( 1 8 5 9 ) in his turn cites both speciesproposedby Mtinster (p. 4 1 , F i g s I 1 4 , I 1 8 , 1 1 9 ) .K e f e r s t e i(n1 8 6 6 p , .1443,PL 130, Fig. 5) only mentions Geoteuthis(Belemnosepia) lata. This is a rather uniform type of shell (Fig. 45); I
Fig. 45.
Geoteuthissimplex (Voltz 1840), drawn from the
original specimendescribedby Miinster ("Geoteuthislata", Beitr. VI, P1.7, Fig. 1) from the Lias e flower Toarcian] of Metzingen;dorsal view, anterior and posteriorparts (beyond the fracturcs)reconstructed.1/3nat. size. m.limit between
carefully reconstructedit from Miinster's original
the central part and the lateral areasof the median plate; s.
material,a task made easy by the growth lines. The
median asymptote;a. lateral asymptote,limiting the narrow
iilustration shows in particular:that the conusvane (c)
laterai plate; .t. lateral line, laterally limiting the "lateral
extendsvery far forwards and tends to extend beyond the junction (at a) with the pro-ostracum;at any rate,
arcuatezone"; c. conusvane,posteriorlybent into a conus.
there is a position (r) Q24 at which the growth lines form a forward-pointing obtuse angle. The lateral
musculature.I have not been able to determinewhether
platesofthe pro-ostracumare very naffow and show a
the insertion of the muscular mantle is limited to the
denselongitudinaland hyperbolic striationbetweenthe two sharply defined asymptotes.The median plate is
the conus vane. Geoteuthis attatnsa size of 30 cm and
tripartite with two lines separatingthe median field,
co-occurswith Belopeltis aalensis in the Lower Lias
which is finely striatedtransverselyand lengthwise,
e fl-ower Toarcian].(At Holzmaden,Paraplesioteuthis
and devoid of a median rib, from the two lateral fields
sagittata lies about 1.3 m abovethat level, accordingto
which show a conspicuouspattemofarcuatestriae.
B. Hauffl. The speciesis widely distributed in the SwabianLias (Frittlingen,Holzmaden,Boll, Ohmden
These shells are almost always associatedwith a large ink sac and often show remains of the mantle
margin or extendson to the dorsal and ventral sidesof
and otherplaces).
76 illustrations,calls the species(l 25) Ommastrephes miinsteriand describesit (1845 [1855],p. a17)in such a way that recognition is impossible. In 1846 he provides an illustration, under the same name, so that its meaningis clear(Pal.Etr.. Pl. 24, Fig. 3, Pal. Univ., P l . 2 1 , F i g . 3 ) . ( T h e a c c o m p a n y i n gf r g u r e 2 , O . cochlearis,apparentlybelongsto Plesioteuthis,cf. p. 1 1 5 ) .W a g n e r( 1 8 6 0 ,p . 4 5 , 6 8 ) c o n s i d e r sG . m i i n s t e r i to be a fragrnentof Tettthopsis (Palaeololigo/ which would have been called Acanthotetrthiscochlearis by Mtinster. I found Mr.inster'soriginal specimenlabeled " Celaeno cochlearis Miinster" in the Munich collections,and I preparedFigure 46a basedon this specimen. The two names have apparently been confusedmany times.
t\
N\I .,'r,\'tl l"'l'i ,'l: ,' ,)k Fig. 46. - Geoteuthismiinsteri(d'Orb. 1846).
G.
The
family
Belopeltidae
Naef
l92l
( S y s t e mp . 5 3 4 ) . Here belong shellsof the kind describedas Belopeltis aalensis(Zieten)(seebelow).
a. Rcconstruction of the she11from old specimensI have seen,especiallythc original specimenof M0nster (Beitr. VII. P1.6. Fig. 1-2), now in the Munich public collections.r/2 nat. size (a-e). r. lateral areas of the median plate; s. curved transverse growth lines; y. lateral asymptote.Locality: Daiting, lithographiclimestones.
The genusBelopeltisVoltz 1840. For the earlier names of the speciesconcerned,see p. 122 on Geoteuthis. Nowadays they are most often
D. Impressionof the animal with the head (/t). arms (fl and
called Geoteuthisbollensis. However, the names of
distinct tracesof the shell in the posteriorpart. v. position of
speciesand genushave a different meaning.(Seeunder
the anterior she1l margin; rz. median groove. Original
B e l o t e u t h i sb o l l e n s i sa n d G e o t e t t t h i s ,
p.
124).
specimenfrom Solnhofen (Hiiberlein collection 1880), now
Belemnosepia andPalaeosepia are redundant
in the Berlin collections.
designationsfor the supposedbelemnitewith which
c. Detailfrom c. enlargedtwice.
these shells were sometimesassociated(p. 109). For
d. Small specimen from Eichstiitt, now in the Munich researchcollections,showing a large ink sac (t) and remains of the muscularmantle (z), originally fiom the Leuchtenberg
Blainville (1825)Loligosepiais a sort of synonymof Sepioteuthis (a Recent squid genus).The following speciescan be consideredas typical:
collection. e. Similar specimenfrom Eichstiitt (in the collectionsof Eichstiitt Lyceum, labelled:"a Geoteuthis")with impression ofthe head.
Belopeltis aalensis (Zieten 1830). This specieswas first describedand illustrated by Z i e t e n ( 1 8 3 0 ,P l . 2 5 , F i g . a , p . 3 a ) u n d e r t h e n a m e Loligo aalensis Schtibler (cf. p. 144) and at the same
Geoteuthismiinsteri (d'Orb. I 8a6).
time also as L. bollensisSchiibler(Figs 5-7). Sincethe
Miinster (Jahrb. 1837, p. 252) apparentlyincluded the
latter designationis used in the samepublication(P1.
original specimenof this speciesin Onvchoteuthis
37, Fig. 1, p. a9) for what we here call Beloteuthis
cochlearis, without a careful description,and later
bollensis. the former name has to be used. ZteIen
figured it without describing and naming it (Beitr. 7,
(1839) used the samenames for Onychoteuthisprisca
P I . 6 , F i g s I a n d 2 ) . D ' O r b i g n y ( 1 8 4 1 ,F e r . & d ' O r b ,
Miinst. 1828(cf. p. 109).The shellsbelonginghereare
introd., p. XL), who received Miinsters data and
the most common onesin Lias e fl-ower Toarcian],but
77 they are always broken and at first sight of diverse
comparativestudy of numerous specimensconvinced
aspect.Hencethe numeroussynonymsand confusions.
me of the correctnessof the first view. The delirnitation
(126) Yoltz (1840) erroneouslydistinguishedthe
of the arcuatelines is enhanced,especially in ventral
speciesBelopeltismarginatus(p. 25, Pl. l, Fig. l), B.
aspect,by a secondaryincrustationwhich lies partly on
s i n u a t t r s( p . 2 8 , P l . 1 , F i g . 2 ) a n d B . r e g u l a r i s .T h e
the arcuate line and produces two narrow, angular
secondform at least belongs here, probably the other
limiting ridges.
onesas well. Mtinster(1843),in additionto Geoteuthis
The medianplate showsno differentiationof lateral
bollensis(Pl. 8, Fig. 1), introducedG. speciosa(Fig. 2)
fields. It is, instead, subdivided into two halves by a
and a G. obconica (P1. 9, Fig. 1), ail of which are
sharp median rib (/r). In ventral view this appearsas a
identical with B. aalen,sis.These "species" reappear
linear band, in dorsal view as a naffow groove flanked
under the name Belemnosepia in the work of
by low elevations,with a very delicate median rib on
d'Orbigny (1845, 1855, p. 440-441); they were
the floor ofthe groove.
illustratedin 1846(Pal.Univ., Pl.28 and 29, Pal. Etr.,
The median plate shows delicate longitudinal lines
PI.25 and 26).(cf.d'Orb. 1850,Prodr.1, p.242).
and arcuategrowth lines. The courseofthe latter varies
Q u e n s t e d t( 1 8 5 8 . p . 2 4 4 ) d i s c a r d s h i s e a r l i e r designationLoligosepiabollensis(1839,p. 163).From
somewhat; sometimes its median part is straight,
1 8 4 9( p . 5 0 8 ,P I . 3 2 ,F i g s 1 1 - 1 3P, l . 3 3 , F i g s l - 5 ) h e
probably unrelated to the growth of the anterior
calls the speciesLoliginites bollensis, before that
margin, but due to variations in the growth of
(1843,p. 252) alsoLoligo bollensis.(Seealso 1858,p.
secondaryincrustations.
2 4 3 , P I . 3 4 ,F i g . l , 1 8 8 5 p , . 5 6 7 ,P l . 3 9 , F i g . 9 . C h e n u ( 1 8 5 9 , p . 4 1, F i g . I l 6 ) m e n t i o n s t h i s s p e c r e sa s BeIopelti s margin atus. Figure 47b shows a reconstructionof the shell,
sometimes slightly concave. Such variations are
The shells are always heavily darnaged. The delicateanteriorand posteriorendsare always missing, so that the conus can only be reconstructedfrom the growth lines: they demonstratea pointed end; the
which allows one to recognizethe charactersof the
relative size of the conus cannot be determined,
species more easily than the fragments which are
however. The curvature of the shell resulted in
generallyavailable:a striking feature is the unusual
separationof the two halves during burial in the Lias
developmentof the conus vane, which extendsalmost
shales.Due to this breakagethe angle between the
to the anteriorend of the shell, the angle in the growth
lateral strips, which would norrrally provide a useful
lines (cf. Geotettthis,Fig. 45, x) at the transitionto the
feature,is alteredand is very variable.Fig. 47b offers a
lateral platesprojecting forwards.Thus a sharpmargin
careful reconstructionof the typical aspectbasedon a
(A) is formed which could be mistaken for the lateral
fine specimen in the Munich collection and study of
asymptote,since it delimits a lateral strip showing a
many others.
pattem of backwardlyconvex growth lines. Figure 47a,
In Germany this speciesis found wherever Lias
however,showsthat the asymptotelies more medially,
e fl-ower Toarcian] is on the surface: Holzmaden,
where the growth lines curve [forwards] (e). The situationof the median asymptotesis also obscuredby
Ohmden,Hondelage(nearBraunschweig).
this metamorphosis,in that the sharp bend in the area of line J is lacking. These "arcuate striae" show a very characteristic
S c h w a r z a c h ,T r i t t l i n g e n , B o l l , M i t t e l g a u , A h l e n , The variants of the aalensis type in the German Lias can hardly be consideredto representdistinct species;at least nothing definite can be said on this
pattern, which may vary slightly. It is much more
question since the state of preservationof specrmens
conspicuousin the ventral aspect,since on the lower
obscures many features that would be useful for
side the shell (hypostracal) layers are scale-like,
tentative distinctions.Examplesfrom French localities
overlapping from anterior to posterior. One might
were not available to me, (I 2 8\ not even as
thereforehave doubts whether the curves ("Ogives" in
illustrations.A11we know about these forms is based
Zieten) of this zone actually representgrowth lines of
on the descriptionsof d'Orbigny and Deslongchamps.
the ostracum,(127) and one may wonder whether they belong wholly to secondary incrustationsa2. A
Related forms from Lyme-Regis (England) are definitely different from the shellsfound in Germany;I
78
J A e
Fig. 47.
Different Belopeltidae;reconstructed from the growth lines.
r/2nat. size.Lias e.
a. Parabelopeltis .flexuosa(Mtinst. 1843) drawn from a specimenin the collectionsof B. Hauff, Holzmaden. b. Belopeltisaalensis(Zieten 1830)drawn from a specimen(markedF) in the Munich researchcollections. c. Loliginites coriaceus(Quenst. 1849) drawn from a specimen(markedE) from Holzmaden(Oppel collection),now in the Munich collections. All drawings were made using the entire availablematerial of the collectionsin Munich, Stuttgart.Tribingen.Holzmaden,Vienna and Berlin. Dorsal view in a-c'.K. median rib (keel); J. median asymptote;e. lateral asymptote;A.lateral lines, limiting the arcuate zones;ir. growth lines of the medianplate;,r-.superimposedtransversestriations;R. limit of the nomally striatedpaft of the vane; O. arcuatelines; H. light zoneof the medianplate. a can reacha sizetwice as largeas the hgure, & and c four times as largc.
have been able to study one specimenin the Ttbingen
shells,in pafticularthe lateralendsofthe concaveparts
collections. This difference was already known to
of the lines do not extendas far forwards.
Voltz (1840,p. 30) who distinguished separatespecies (morethan necessary!). Buckland(1836,Pl. 28, Fig. 6,
Following Voltz (1840)we distinguishat leastone morespeciesol Belopt,ltis
PL.29, Fig. 2) mentions only one form, similar to the German shells,as "Loligo" aalensis,while leaving all
B elopeltis b uckl ancli Y oltz.
the other ones, which are more clearly different,
Here belongs the material described by Buckland
unnamed.A very characteristicfeatureis the curvature
( 1 8 3 6 ) ,P l . 3 0 , P l . 2 9 , F i g s 1 a n d 3 ) . T r e a t e db y V o l t z
of the concave,arcuateparts of the growth lines. lt is
( 1 8 a 0 ,p 3 0 ) a s B . b u c k l a n d(iP 1 . 3 0 )a n d B .a c u m i n a t u s
shallowerin Buckland'sspecimens than in the German
(P1.29,Fig. 1), and as an undeterminedspeciesin Pl.
79 2 9 , F i g . 3 . ( c f . d ' O r b i g n y ,1 8 4 6 ,P a l .U n i v . ,P l . 2 9 , F i g . i : P a l .E t r . .P l .2 6 . F i g .3 . ) .
Loliginites coriaceus Quenst. I 849 (: TeudopsisagassiziiDeslongchamps1835). The first satisfactorycharacterizationof this speciesis
The genusParabelopeltisNaef 1921 (Systernp. 534).
d u e t o Q u e n s t e d(t1 8 4 9 ,p . 5 1 2 , P l . 3 4 , F i g . 5 - 8 ) .T h i s source is therefore definitive. However" Mtnster's
Only one speciesbelongshere:
(1843, Pl. 6, Fig. 6) Sepialitesstriatulus probably belongs here as well. Without any doubt "Teudopsis"
P. Jlexuosa(Mtinster 1843).
D e s l o n g c h a m p( 1s 8 3 5 ,p . 7 2 , P L . 5 , F i g . 1 5 )b e l o n g st o
Original designation:Geoteuthis.flexuosaMtinst. (VI),
this species; thus that problematic genus can be
Pl. 9, Fig. 2. Rarely taken into account,often confused
discarded.Although Deslongchampsdid not provide a
wtth Belopeltis
A c c o r d i n g t o W a g n e r ( 1 8 6 0 , p . 6 2 ) i t b e l o n g st o
clear-cut illustration or description, Quenstedt establishedthe identity with his original species.He
Geoteuthissagittata; the overall aspect of poorly
reports(1858,p. 245):
aalensis and Geoteuthis ,simplex.
preseled specimensindeedresemblesthis species.
"Study of the original specimens in the fine
D'Orbigny (1845, 1855,p. 431) lists the speciesas
collectionof Mr Deslongchampsin Caenconvincedme
Belemnosepia .flexuosaand gives an illustration(Pl. 31, F i g . 2 ) . L i k e w i s ei n 1 8 4 6( P a l .U n i v . ,P l . 2 5 , F i g . 2 a n d Pl. 26, Fig.2. - Pal. Etr., P|.22, Fig. 2 and Pl. 23, Fig.
that Tettdopsisagassizii is identical with cctriacetrs,a description.Therelorethe name I proposed,althoughit
2 ) . C h e n u( 1 8 5 9 ,p . 4 1 , F i g . 1 1 5 ,1 2 0 )l i s t si t u n d e rt h e
is younger,must be conserved"(p.245). (cf. below p.
samename.
r44).
This speciesmainly occurs near Holzmaden in the Upper Lias efl-ower Toarcian]. I have also seen
fact that could not be decided from the drawing and
Here probably also belongs: Sepialitesstriatulus
identifiable specimensin the Braunschweigcollections
Q u e n s t e d(t1 8 a 9 ,p . 5 1 5 , P l . 3 6 , F i g . 2 0 ) . D ' O r b i g n y (1845, 1855,p. a31) lists the speciesas Belemnosepia
(129) (from Hondelagenear Braunschweig).(Same
a g a s s i z i i a c c o r d i n g t o D e s l o n g c h a m p s ,w i t h o u t
age). Although in these areasit seemsto be younger
mentioning any characteristicfeatures.The illustration
than Geotetrthis simplex and Belopeltis aalensi,s,it
(p. 31, Fig. 3) is a totally false reconstructionwhich
clearly offers an intermediate form between the two
contains no characteristic feature other than the
types(Figs 45 and47b).
longitudinalstriation(cf. 1846,Pal. Univ., p. 25, fg. 3;
median rib: this rib (in dorsal view) forms a narrow
P a l . 6 t r . ,p . 2 2 , f g . 3 ) . Q u e n s t e d(t1 8 5 8 ,p . 2 4 4 , p . 3 4 , fg.8, and1885p , . 5 0 7 , p l 3 9 , f g . 1 1 ) l i s t st h e s p e c i e s
ridge between two narrow grooves. On either side of
w i t h i t s v a l i d n a m e .( 1 3 1 ) C h e w ( 1 8 5 9 ,p . 4 1 , f g . 1 1 3 )
these are reinforced plates. The rnedian asymptotes
still usesthe nameBeletnnostpia agassizii.
form (130) conspicuousfree margins lat the anterior end] and the growth lines curve forwards only weakly
The particular similarity to the other two generaof this
laterally, forming a rounded angle [in contrast to the
is preserved,without the broken tip. In theseexamples
sharp point in Belopeltisl. The formation of the
the sharp longitudinal striation and the densely set,
characteristicarcuate lines is seen in its early stages
very distinct growth lines of the conus vane are
here. On the inner side they are still as usual bounded
conspicuous;they suggesta reconstructionsimilar to
by the rnedian asymptotes.The median plate shows a
Geoteuthis.Moreover long specimens(up to 30 cm)
pronounced longitudinal striation and a (fairly
are known in which the anteriorend has only tracesof
inconspicuous)demarcation of lateral fields with
the longitudinal striation, further obscuredby breaks
It is distinguishedfrom the previous speciesby its
different,generallydarkercoloration.
family is not striking: quite often only the posteriorend
and deformation.The conusvane extendsfar forwards but shows quite different features: instead of the
The genusLoliginites Quenstedt1849
longitudinal striation, which (apparently)is lacking
(s. restr.Naef 1921).(Systemp.52\.
altogether,there is a very fine transversestriation on the exceedingly delicate layer, below which the
The follorving speciesbelongshere:
muscular mantle extendedmedially. Only particularly well preservedspecimensshow the relationshipof the
80 and a delicate anterior lateral part, the latter having apparently been formed only at later stages, superimposed on the muscularmantle which retained the site of its inserlion. Affiliation with Belopehls appearslikely given the forward extension of the conus vane and the overall form. But the typical arcuatestrip patternof Belopeltis seemsto be lacking. It is indeednever observedin such a conspicuousdevelopment.However, consideringthat the lateral plate cannot have vanished between the medianplate and the conusvane! one may examinethe lateral parts of the apparentmedianplate more closely, and then one will find this: a sharply demarcatedzone correspondingto the zone of arcuatelines shows r:ery delicate transversestriae crossing the longitudinal striations; these delicate transversestriae follow the same course as in Belopel/is. This is particularly distinct in the central to posterior pafts of the shell, if the latter is well preserved.For this observationa magnifying lens is often necessary,but then one will always find this pattern. (In Fig. 4lc, at o, the lines have beenunnaturally enhanced). The posteriorpart of the medianplate in most cases shows a conspicuouslight zone and darker lateral fields. The median (132) line is occupiedby a delicate keel which tapers anteriorly. Other longitudinal lines are variable and virtually disappearanteriorly so they cannotbe includedin an unambiguousdescription. This speciesis common in the Upper Lias e fl-ower Loliginites zitteli, from the original, rL nat. size.
Toarcian]. I have seen specimensfrom Trittlingen,
The figure emphasizesonly the visible parts of the animal,
Holzmaden,Mdssingen(in Swabia) and from Curcy
without any further interpretation.6. arm stumps; z. lower
and Trois-Monts (France, Calvados). The gladius
mandible;a. eye (!); k. cephaliccartilage;1.nuchalfold (?);
attains a length of about 40 cm. According to
l. supposedposition of the funnel; ,t. ostensible funnel
Q u e n s t e d (t 1 8 4 9 , p . 5 1 2 - 5 1 5 )i t s o m e t i m e se x h i b i t s well preserved remains of the soft body; but this
Fig. 48.
(accordingto Fraas),perhapsthe anterior end of the liver, as a darkly stained steinkern, perhaps the mud filling of the funnel cavity; l. longitudinally striated part of the mantle, probably an integumentalstructure;n. transverselystriated muscular mantle; L ink sac; /. gap in the muscular mantle (indicating the shell which has totally disappeared)l.l fin musculature.- A funnel notch is not recognizable;the pafi of
outstandingobserverdid not have sufficient special knowledge to interpret them. The stomachwith scales and bones of Leptolepis, and remains of the muscular mantleand ink saccan be recognizedwith certainty.
the mantle missing on one slab is visible on the countelpaft. The cephalic cartilage is said to show a muscular structure
H. The family Lioteuthidae nov. fam.
(Fraas, p. 223). Of course such a fine distinction of anatomicalelementsis imoossible.
Herebelongs: The genusLioteathis nov. gen.
two sorts of fragments. It can then be seen that the conusvane is subdividedinto a robustposteriormedian
With a singlespecies:
=
81
.t
3 Lt
5
c.
Fig.49.
Teuthoidshe1lsfrom the Cretaceous.
of Bohemta. minorafterFritsch(1910,Pl. 16.Fig. 13a).'/r nat.size.FromtheUpperCretaceous a. Glyphiteuthis b. Trachytufihislibanotica (Fraas1878).Drawn from the original specimendescribedby Fraas(1895, Pl. 6, Fig. 3), now in Stuttgart, and two larger specimensfrom the Berlin collections.The proportionscorrespondto one ofthe latter (purchasedin 1877, 1/1 fomerly ownedby Hakel), drawn nat. size.The other specimensare from Sahcl-Alma(Upper Cretaceousof Lebanon). t/5 c. "sepialitessohel-almae" (O. Fraas 1878.MS) from the samearea!spccimenin the Stuttgartcollections.Sketch nat. size. One can recognizethe mantle sac (4), stomach(5), ink sac (3) and rcmains of the shell, especiallytwo lateral ribs (2) of the median plate (cf. Fig. 47b), and the head(1). The overall aspectisthat of Geoteuthis.
Lioteuthis problematica n. sp.
simple fom, which could be mistakenfor a belemnite
The peculiar featuresof this type require creation of a
pro-ostracum if it were not for those peculiar lateral
new family, which is rather isolated.fNote: there have
plates.
been severalrecentattemptsto restudyand characterize this unsatisfactorygenus]. Here I place a gladius from the Lias (Fig. 41d),
I. Doubtful forms of Prototeuthoidea.
which I found in the Tiibingen museum (reg. Nr. 8764). It cannot be united with any other form and
l. "Loliginites (Geoteuthis)zittuff' E. Fraas1882.
must therefore be placed in a separatefamily and
HerebelongsFigure48.
genus. Unfortunately, however, it is rather poorly
The original specimenis housedin the natural history
preserved;this of coursehamperscharacterization of
collections at Stuttgart; it is an exceptionallywell
thesetaxa. One can vaguely distinguish:median and
preserved complete animal, which (133l became
lateral piates, the former with fine longitudinal folds
separatedinto a dorsal and ventral part when the
and a distinct,weak medianrib in its posteriorpart, the
calcareousconcretioncontaining itar was split in two,
latter with very peculiar growth lines. Roughly in the
so that both surfacesprovide almost identical pictures
middle lies a moderatelysized ink sac. The conusvane
of the animal in dorsal and ventral aspect.However, closer inspectionrevealedthat the (fide Fraas,p. 223)
and conus are no longer recognizable.It is a curiously
82 2. "Sepialitessahil-almae"O. Fraas1878(MS.). Here I place a fossil from Lebanonnamedby O. Fraas, now housed in the natural history collections at Stuttgaft.The label reads:"SepialitesSdhil-AlmaeFrs. Sahil-Alma Ecire 1878". It is an animal showing an overall resemblanceIo Geoteuthis(Loliginites) zitteli, without distinct remains of arms but with clear traces of viscera and shell. The head and mantle sac are only vaguely outlined, the moderatelysized ink sac lies far anteriorly. Of the shell one recognizes(in this as well as in a similar but less well preservedspecimen)the outline of the pointed triangular median plate (135), Fig. 50. - Arm crown of a teuthoid fiom the Cretaceousof Lebanon. "Plesioteuthisfraasi", probably identical with "Dorateuthissyriaca" (p. I l8) from Sahel-Almaaccordingto W o o d w a r d( 1 8 9 6 ,p . 2 3 3 , P l . 1 ) . ( c f . O . F r a a sl 8 7 8 , " F r o m thc orient". p. 90). Three pairs of arms differing in length.
which however is deformed.It seemsto be delimited by two thick, curved ridges or rods, u.hich perhapscan be comparedto the peculiar arcuatelines of Belopeltis aalensis.(Seerough sketchof Fig. 49c).
plus one longer arm. traces of suckers or brachial cirri (supports of protective membranes),and the mandibles of
3. Ommatostrephes meyrati Ooster1857.
typical form are recognizable.There is nothing strikingly I special. l2 nat. size.
A doubtful fossil; it was interpretedby Ooster(1857; cf. 1863,p. 5, Pl. I, Fig. l-2) as the conus of an "Ommatostrephes" sensttd'Orbigny (p. I 15). It is from the Neocomian of Rufisgraben near Beatenberg
"beautifully preservedeyes" were painted on the stone
(Switzerland,Canton Berne); it does not show any
a n d i n d e e d r e p r e s e n ta n e r r o n e o u si n t e r p r e t a t i o n
morphologicallynoteworlhy or characteristictraits of a
attemptedby the preparator(?). The overall aspectis
true teuthoid.
essentiallygood, as well depictedby Fraas' original figure, rather than characteristicdetails. There is no distinct trace of the shell, hence no possibility of
K. The MesoteuthoideaNaef 1921
making comparisonswith known shellsor types.
(Cephalopoda, vol. I, p. 145).
The overall aspect,at least, is that of Belopeltis aalensis, and the striking size of the ink sac can be
Diagnosis: Mesoteuthoidsare fossil teuthoids(Lias
viewed as a characteristicfeature of a prototeuthoid relatedto this form. As to the details.however.I cannot
[Lower Jurassic]to Cretaceous)in which the anterior margin of the median plate of the gladius forms a
subscribeto the reconstructionproposedby Fraas.The
nalrow cur-ve,a point or a pointed process,[the median
"funnel" is probably the anterior end of the liver, the
plate isl folded along the midline to form a broad gutter
eyes are only hinted at by some traces lying further anteriorly, the arms are only representedby stumps
which is open ventrally ("kee1")aa,and without asymptotessharply demarcatingthe lateral plates,- rn
(eatenoff or rotted away) so that nothing can be stated
which the conus vane is leaf-shapedand occupies
about arm length. In contrast,(134) the "wings" of the lower mandible are clearly recognizable,indicating the
about one halfofthe gladiuslength,gradingposteriorly into a spoon- or ladle-shapedconus,- in which lateral
position of the buccal mass, as are the lateral parts of
asymptotesare defined by a slight concavity of the
the cephaliccartilage.See Figure 48, which is easily
growth lines.
comparedwith Fraas' original illustrations.There are
These forms apparently lead, through a series of
well preservedmuscular structuresof arms, mantle, and - as far as I can see- remainsof terminal fins. The
transitions,to the recentLoligo-hke shells(136) which
illustrations of these structuresas provided by Fraas
the following trends: |. The calciJicationof the gladius
indicatea strong similarity of the tissuesto those of
decreases, the shell becomeshorny, thin and apparently
recentdecaoods.
flexible. 2. The anterior end o.f the median plate
are united in our Metateuthoidea.This is expressedby
=
83 :'tt'omesnarrower and, in extreme forms, drawn out :ntoa long pointedprocess.3. This modificationentails t rtarrowing of the v,hole medianp/ate to the advantage .,i the lateral plates.The median plate finally becomes .r narrow strip which forms a median rib called the ''rachis" betweenthe leaf-shaped lateralplates.4. The 'ttt€ral plates become increasingly limited to the :,rtsteriorpaft of the shell, taperingoff anteriorly along the rachis,the anteriorparl of which thus becomesfree, iike the stalk of a leaf. 5. The narrowingof the rachis is partly causedby its longitudinalflexure so that it forms .r dorsal "keel" while the lower side shows a broad groove. 6. The conus becomes rounded, ladle- or \poon-shaped. The transformationof the broad medianplate into a tl'eerachis,seenespeciallyin the family Beloteuthidae, conditions anatomical changes in the area of the nuchal cartilage, the head/foot retractors and the stcllate ganglia, as shown by a comparisonof Figures -10 and 58. The gradual modifications thus gain a certain significance,justifying the separationof the prcsentgroup. Voltz (1836) alreadyreflectedon the rnorphologicalcharacterof these forms. He writes, rritlr regard to a specific genus(Beloteuthis),which he calls "Tetrdopsis" (p.3): "One can thus define Tetrdopsis as follows: sheath lacking, alveolus very incomplete,horny; dorsal region (our median plate) vcry elongate;hyperbolar regions (lateral plates) very rvell developed,especiallyin their lower or posterior
Fig. 51.
part, no ventral region and no chambers".
This
reconstructed from the original specimen from Eichstiitt,
characterizationis evidently unsatisfactoryin part, for
describedby Zittcl (1885, p. 516; cf. Grundztige1921,p. 595), in the Munich teachingcollections(no. 34). 1/1nat. size.-.r,. only slightly markedmidline with anteriorlyconvex
rvantof clearconceptsof form.
T r a c h y r e u t h i s h a s t i . f o r m i , r( R i i p p . 1 8 2 8 ) ,
growth lines; :. tubercular zone of the median plate.
L. The family Trachyteuthidae Naef 1921 (Systemp. 534).
reminiscentof Sepio;sp. lateralplate; .r. uncalcifiedmarginal zone;.ra. lateral asymptote;.sb.lateral arcuatezone; v. area with superimposedlines concavebackwards,belongingto the conus vane along with sb and cf.
D i a g n o s i s : S t o c k y m e s o t e u t h o i d sw i t h s t r o n g l y calcified,more or less cuttlebone-likeshells,showing knobby granulationsin the middle zone (137) of the
similar to those of living Sepia, are neverthelessso
dorsal side. Typical species:Trachyteuthishastilbrmis
clearly different that they have to be placed in a
(Riipp.).(SeeZittel,Grundziige,1921,p. 595).
different genus, which I
caII Trachyteuthis.The
collectionsat Ansbach contain excellent shells of two The genusTrachyteuthisH. v. Meyer 1846.
species which I designate as L
oblonga and T.
ensi.fbrmis, respectively;a fragment illustrated by H. v. Meyer (18a6,p. 598) characterizes this form as a
Mtinster (VII, Pl. 9, Fig. 3) appearsto belong to the
distinct genus, different from Sepia (in contrast to
latter; the former speciesis not mentionedin Mtinster's
generalideas of his time): "The shells,which are very
notebooks.On the lower surface of these shells I
84
Fig. 52. - Reconstructionof Trachyteuthis hastiforrrls drawn from the well preservedspecimenfigured by Crick (1866, P1. 14, Geol. Mag.). The tentaculararms,the suckers,the protective membranes,and the eyeshave been addedin their typical forms. The fins have been addedfrom a different specimen(where they are perfectlypreserved)ftom Eichstdtt(specimenin the Eichstiitt t12 Lyceum). nat. size..s.lateral margin of pro-ostracum;f fin base (distinct impressionof articular cartilage);lz. muscular mantle.
noticedpeculiar,small,round impressions,sometimes
illustrations of Mrinster's "species" and withdraws
with radiating structures or angular depressions,
some of them, maintainingthe following species:S.
vaguely reminiscent of the articular surfacesof stem ossiclesin certain crinoids." - He "found that they
lingtrata, obscura, regularis and gracilis. S. venttsta Miinster 1837,later d'Orb. 1845, 1846,Pl. 7 doesnot
form a seriescorrespondingto the median line of the
belong here. See under Argonautat):
snells
hastiJbrmisand ensiformlsMeyer 1846,p. 598; 1856, p. 106,Pl. 19 : S. hastifurmisQuenstedt1849,Pl. 31,
The shellsbelonging here in fact representa single species,which hasalreadybeenwell described:
Trachytettthis
Fig. 25, Pl. 32, Fig. 1, p. 493. Coccoteuthis latipinnis Owen (1855,p.124, Pl. 7) probablyalsobelongshere.
7. Trachyteuthis hast{ormis (Rrippell 1829). This species is already illustrated in
MoreoverseeFraas(1855,p. 86); Wagner(1860,p.
Knorr
6-18; Chenu (1859,p. 45, Figs 132-137).(Here again
(MerkwtirdigkeitenI, Pl. 22, Fig. 2), but only Rrippell
are the speciesof Mtnster and d'Orbigny); Keferstein
describedit carefully(1829,p. 9, Pl. 3, Fig. 2). (138) It
( 1 8 6 6 , p . 1 4 4 1 ,P l . 1 3 0 , F i g . 1 0 a s " C o c c o t e t r t h i s "
is a rather abundantform in the Solnhofenlimestones,
hastiformis;Fischer(1887,p. 357).(Fischerseesherea
but it is never perfectly preserved.Different fossils
transitionfrom the Sepiaphorato the Chondrophora).
therefore present a variable picture: they often show
Trachytetrthisattainsunusual sizes (Wagner 1860,
only isolated layers or impressionson a slab, or they
p. 1l): specimenswith gladiuslength up to 75 cm have
are deformedand laterally compressed,flattenedinto a
been reported once, others with 40-50 cm more than
singleplane,etc.Hencethe numeroussynonyms. Sepia hasti/brmls Rrippell \829 : S. antiqua,
have measuredup to 1.5m in total length.
caudata, linguata, obscura, regularis, gracilis Miinst. 1 8 3 7 .( S e ed ' O r b i g n y1 8 4 5a n d 1 8 4 6 ,P l . 5 - 6 ; h e g i v e s
once. Since the arms are very long, the animals must This species occurs widely in lithographic limestones:Nusplingen, Solnhofen,Eichstiitt, Daiting.
85 Fine specimensare housed in the Munich collections;
counterpart,I was indeed surprised to find such far-
isolatedarm crowns were availableto me in Berlin.
reaching similarities. The following detail is particularly striking: as in Sepia, the muscular mantle
they correspondexactly with the illustration by Crick I 1896)and with partsobservedon the animal.
extendsto the inner side of the "wings", i.e. the conus
The shape of the shell is oval in the smaller posterior half, broadenedby the "wing" of the conus
the prototeuthoids,in contrast with the metateuthoids
vanes, very slightly pointed at the anterior end, thus
(Fig. 58). A similar life style can probably be assumed:
emphasizingits mesoteuthoidcharacter.The slightly
the stocky overall aspect and the weight of the shell
pointed end, which is not recognizable in poorly
suggesta flore or less pronouncedbenthic variant of
preservedspecimens,grades into a very flat, narrow,
the teuthoidtypea5.
barely visible median keel. There is no longer any
vane(seeFig. 35c) and thus placesthis genuscloserto
Fischer(1887, p. 357) seeshere a transitionfrom
naturaldemarcationbetweenmedian and lateralplates; they are both marked- in additionto the growth lines -
the "chondrophores", i.e. teuthoids, to "sepiophores".
by delicate longitudinal iines which convergetowards the conus(Fig. 51).
2. Truchyteuthislibanotica (O. Fraas1878).
(139) The lateral asymptote (sa) is clearly recognizable, though ratherincor.rspicuous; it delimits
the
This specieshas been described and illustrated as "Geoteuthis"libanotica(p. 90) by O. Fraas(1878);it is
the conus vane. The latter comprises three fields (labelledsb, v and cf in Fig. 51), the medianone being
said to have eight "tuft-like" arms, and hence to be
easily comparableto the correspondingone in Fig. 73a,
crown. I examinedthe original specimensin the natural
45 and 47. The adjacent field shows a retrograde
history collections at Stuttgart and in the Berlin
transversestriation which crossesthe growth lines; it
Museum of Natural History, and I made Figure 49b by
probablyis relatedto the hn inserlion.
combining them. Z. libanotica appearsto be a small
The conus is rounded,spoon-shaped, as in all other
similar to the previous specieseven as regardsthe arm
mesoteuthoids;it is totally blunt, in contrastto the
form; the pro-ostracum is narrower than that of the preceding speciesand thus suggestsa less marked
metateuthoids, which in this respectsometimeslook
differencefrom the Beloteuthidae.
rnore like prototeuthoids.Particularly striking is the surface granrilation of the central field of the dorsal
The genusGlyphiteuthisReuss1854a6.
side, which is very reminiscent of Sepia (e.g. ofticinalis).This granulationis made up of round knobs
Two speciesfrom the Bohemian Cretaceousbelong
which look like the convex part of a pressbutton; they
here, included in Trachyteuthisby Fischer(1S87,p.
rnay have servedfor the attachmentof the dorsal skin.
357).
Another striking similarity is the delicate,uncalcified marginal zone (n) which is particularlywell seenin the well preservedfragmenton which this part of Fig. 5l was based. Of course this feature is common to all
1. Glyphileuthis ornata Renss (cf. Gl. ornataFritsch1910,p. l3-14, Pl. 5, Fig. 6). From the marly lirnestone, the "Pliiner vom
c a l c i f i e ds h e l l s (. S e ee . g .F i g s 3 5 a ,b , 3 7 c , 3 8 g , f , 9 0 ) .
Weissen Berge" near Prague. A well preserved
Flrrthermore considering the overall shape, one can
impressionof the dorsal side. The spatularshell is 23.3
understandwhy earlier authorsclassifiedthis genus
cm in length, the anterior end with a blunt point,
with the sepiids.
posteriorly broadening,then with a conus vane and an
(110) This resemblanceis all the rnore striking as it
oval spoon-like end. (111) The spoon is only 8.2 cm
extends to the soft parts (Fig. 52). I was able to
long and 7.76 cm wide, the posterior part of the pro-
complete my reconstruction using the beautiful
ostracumis 5.46 cm wide. The strongmediankeel is
specimendescribedby Crick (1896), which is now housedin the collectionof O. Abel (Vienna) fnow in
4.5 mm wide, most stronglydevelopedfrom the middle part onward. On either side it shows many transversal
the Natural History Museum, London], and a fine
incisions with interveningelevations.In the middle
illustration, rnadein Eichstritt,of a specimenshowing
paft there are 9, at the ends 12 incisionsper (Viennese)
both fins impressed on both the slab and its
inch. Each incision bears three protuberanceson its
86 upper margin; the middle protuberancebeing the
which have been adequatelydescribed.Theseshellsare
strongest. On either side of the keel lies a granular
particularly abundantin the Lias e fl-ower Toarcian] of
field; it is broadestin the middle part where the largest,
Swabia.
pointed tubercles are connectedin irregular rows. Otherwisethe shell is smooth, only locally showing
The genusBeloteuthisasMtinster I 843.
lines that lie parallel to the aperture.Remains of soft pafis are lacking. The following speciescould be a juvenile form of Cl. ornata.tcf. Fig. 49a;.
Here belong part of the shells named Teudopsisby (1835) (cf. p. 130),for which Fischer Deslongchamps (1887, p. 353) used the correctedname Teuthopsis, including Teuthopsis Wagner (p. A1). Teudopsis tn
2. Glyphiteuthisminor Fritsch 187247
part belongs to Loliginites Quenst. s. restr., and Belotetrthis has long been establishedas a generic
( l o c .c i t . ,p . 1 7 ,P l . 1 6 ,F i g . 1 3 ) . Found in the yellow "Baupliiner" near Lipenec (Lipenz) near Laun. Length 16 mm, width 12 mm. Spoon-shaped,anterior part broken, apparentlywith a pointed end. On either side of the mediankeel lies a rounded ridge. No granulation as in G. ornata. The length of the conus vane is two fifths of the total length; but one has to allow for the anteriorend which is missing.
n a m eb y M i i n s t e r( 1 8 4 3 ,P l . 5 , F i g . 1 ) ( F i g . 5 3 b ) .A s long as really distinctive generic charactersare not known, the diagnosisis the sameas for the family. A distinctive feature of the beloteuthidscompared with recent squids must have been the Sepia-like stockiness.(However, there are much more slender sepiids and almost equally short speciesof squid-like forms, as alreadynoted by d'Orbigny 1845, 1855,p.
Evidently these are Trachyteuthis-likeshellswhich show a clear transitionto the Beloteuthidae.One could imagine a seriesTrachyteuthishastformis, libanotica, Gll,phiteuthis ornata, minor. But all these forms are younger than the Beloteuthidae,which were already
336, who citedLoligo brevls Blainv.). It is striking that these shells almost always lack an ink sac and other soft parts; the latter were probably rapidly separated from the shells, or the shells were buried only after destructionofthe soft oarts.
thriving in the Lias period and which led directly to the recentteuthoids. Beloteuthis subcostataMiinst. I 843. Here belong:Loligo nov. sp. Quenstedt(1843,p.254). M. The family BeloteuthidaeNaef 1921
B e l o t e u t h i s s t r b c o s t a t a( i b i d . , P l . 5 , F i g . 3 ) . B .
(Systemp. 535).
Jusiformis(ibid. MS). B. subcostatad'Orb. (1846,Pal. U n i v . , P l . 1 6 , F i g . l - 2 , P a l .E t r . , P l . 1 3 , F i g . 1 - 2 ) .
A number of shells belong here, apparentlyrelated to
Loliginites subcostatusQuenst.(1849,Pl. 32, Figs 7,
Trachyteuthis, and showing a striking similarity to
8). (143) L. giganteus(ibid.: specimensmeasuringup
recentsquids,a fact that has been noted ever sincethey
to 50 cm in lengthl). Beloteuthissubco.stata d'Orb.
were discovered,so that even d'Orbigny (1845, 1855)
(1850, I, p. 2al). Here are also placedB. substriata
placed one speciesin the genusLoligo (p. 336) and the
Mtinst. (with good reasons)as well as B. acuta and B.
remainingonesat leastin his "Loligidae".
venusta (without any good reason)(cf. Miinster 1843,
(142) Diagnosis'. Beloteuthids are stocky mesoteuthoidswith leaf-shapedshells which from a regular pattern of growth lines
apart
were smooth,
Pl. 6, Figs 4 and 5, and Pl. 14, Fig. 2, and my Figure 54). TeudopsissismondaeBellardi 1856 (cf. Sismonda 1855,p. 1195).BeloteuthissubstrialasChenu(1859.p.
glossy and only weakly calcified, showing a strong
38, Fig. 103,B. subcostatus, ibid. Figs 104, 106, 107).
mediankeel and ending anteriorlyin a blunt, distinct or
B. strbcostattts Keferstein1866 (p. 1,443,Pl.130,Fig.
slightly protractedtip.
9). B. leckensbyiTate and Blake 1876 (p. 314, Fig. 2,
A typical speciesrs Beloteuthis bollensis ZieLen. Relatives are known from the Lias and from the Cretaceous.We unite them in the samegenussinceany differentiation would seem artificial for those forms
P l . 4 ) . B . s u b c o s t a t aS t r o m e rv . R . 1 9 0 7 ,p . 2 5 1 , F i g . "'-This species(Fig. 53a) is strongly reminiscentof Trachyteuthis,but the shells are smooth and glossy,
87 Fig. 53.
She1lsof Beloteuthis from the
Lias e flower Toarcian] of Swabia. t/2 nat. size.Dorsalviews. u . B e l o t e u t h i ss u b c o s t o t aM i i n s t . 1 8 4 3 (after the original specimen dcscribed in Beitr. VI. Pl. 5, Fig. 2; in the Munich public
,/
riir
collections) from Ohmden. Largest form, attaininga lcngth of r/, m. b. Beloteuthisbollensis(Zieten 1830, Pl. 37). (From specimenno.281 in the Munich teaching collections; original specimen describedby Miinster VI, Pl. 5, Fig. 1, "8. tmpullaris"). cf. Zitte\ (1885, p. 518, Fig. 7 t2). c'. Cross section of a specimen(Munich) prescrved three-dimensionally in bituminous limestone from Holzmaden (Mrinst. VI, Pl. 6, Fig. l). (cf. Quenstedt 1 8 4 9 ,P 1 .3 2 , F i g . l 5 a n d Z i t t e l 1 9 2 1 ,p . 5q5l d. Lateral view reconstructedfrom similar specimens;curvature of the posterior end liom a specimen liom the Lias e flower Toarcian]of Curcy (Calvados).houscd in 'ia Stuttgart. nat. size. ,r: marks the lateral arcuatezone.
especiallyon the dorsal side, and the shapeis broader,
growth lines are incorrectin the anteriorpart. They are
drawn out anteriorly into a pointed arch. As in
parallelto the margin.
Trachyteuthis and Geoteuthis, the median part of the
Here also belong: Teudop,sisbollensis (Zieten)
conus vane is differentiatedas the "lateral arcuate
Voltz 1836,p. 6, 1840,p. 33. Loligo
strip" (r) by a bend in the sharplymarked growth lines. From the posterior pole thin longitudinal lines radiate
Quenstedt1843, p. 254. Beloteuthisampullarls Miinst. 1 8 4 3 ,P I . 5 , F i g . l , P I . 6 , F i g . 1 . B e l o t e u t h issu b s t r i a t a
across the growth lines; the longitudinal lines in
M i i n s t . 1 8 4 3 ,P l . 6 , F i g . 5 ( ? ) .B . v e n u s t aM i i n s t . 1 8 4 3 ,
particular characterizethe area of the median plate
Pl. 14, Fig. 2 (cf. p. 138).Sepialites gracilis Mtinst.
lying on either side of the strong keel. As in other
1843,Pl. 14, Fig. 5 (?). Teudopsisampullarls d'Orb.
schiibleri
Beloteuthidaethe growth lines are (144) regular, but
1 8 4 5 ,P a l . U n i v . p . 1 9 0 , P l . 1 1 , F i g . 1 - 2 . T e u d o p s i s
particularly sharpand ratherwidely spaced,like waves
a m p u l l a r i s d ' O r b . 1 8 4 5 .P a l . E t r . , P l . 8 , F i g . 1 - 2 .
following one another.
Teudopsis bollensisd'Orb. 1845,Pai. Err., Pl. 11, Fig.
This form attainslarge sizes,up to 50 cm (Quenst.
3. Loliginites schtibleri Quenstedt1849,p. 499, Pl. 32,
1858,p. 506); it occursin Lias e fl-ower Toarcian]of
F i g s1 4 , 1 5 ( c f . 1 8 5 8p , . 2 4 3 , P l . 3 4 ,F i g . 9 , a n d 1 8 8 5 ,
Swabianear Ohmden,Holzmadenetc., and in England
p. 505, Pl. 39, Fig. a). Beloteuthis ampullaris,
and southemFrance.
s c h t i b l e r ib, o l l e n s i sC h e n u 1 8 5 9( p . 3 8 - 3 9 ,F i g s 1 0 1 , 1 0 2 , 1 0 5 ) .T e u d o p s i sc u s p i d a t aS i m p s o n1 8 5 5 ,p . 2 1 .
Beloteuthis bollensis (Zieten I 830).
Ibid. TateandBlake 1876,p. 314,Pl. 4, Fig. 3.
Loligo bollensis(Schtibler)Zieten 1830,p. 49, Pl. 37,
This species is even more common5othan the
Fig. lae.Ostensiblyan,,Appendixto Pl. XXV, Fig. 5,
precedingone; given the more advancednarrowing of
p. 34" (which in fact belongsto Belopeltis aalensis;q.
the pro-ostracum it is placed after the preceding
v.). The figure is characteristic(outline, keel), but the
species.The pro-ostracum is nearly straight, with an
88 acute anterior margin. The growth lines and
p. 500) who then concluded that Teudopsis bunellii
longitudinal lines are less conspicuous and more
must be identical with Loliginites schiiblerl (our
densely set. In some specimensthe posterior end is
Belotettthisbollensis).Indeed,Quenstedtnoted that all
well preserved in its three-dimensionalcurvature,
the specimensof Deslongchampsbelong to Lias e
embeddedin bituminous limestone rather than slates. Thereforethe shell can be completelvreconstructed
fl-ower Toarcian] rather than to the "grand oolithe" (Middle Jurassic)as assumedby d'Orbigny (1841, p.
(Fig. s3d).
19, 1850, 1852);this meansthat stratigraphically they
The speciesis smallerthan the precedingone; it has a nearly identical distribution.Large, well preserved
are very closely related to the specimens of the
specimens,when reconstructed,measureup to 20 cm
Swabian and Franconian Jurassic. (See Quenstedt 1885,p. 505).The specimenin questionin fact showsa
in length.(SeealsoDawkins1864,andbelowp. 161).
nearly complete morphological similarity (146) to
(115) The following species, which has been
Beloteuthis bollensis,showing especiallythe spoon-
considered as distinctdue to the lack ofprecisionofthe
shaped curved posterior end of this species. The
original figures and the arbitrary separation by
original specimensof Deslongchampsalso show an ink
d'Orbigny (1845,Pl. 20), is likely to be identicalwith
sac, something only rarely observed in ,8. bollensis.
Belotettthis boIlensis:
Thereforethe following, relatedspecieshas to be taken into consideration.
Beloteuthisbunellii Deslongchamps1835. Accordingto Quenstedt(1885,p. 505) this speciesalso
B eloteuthis acuta Mrjnst. 1843.
belongs to the Lias e fl-ower Toarcian] rather than to
Here belongs the shell form well figured by Mtinster
the Middle Jurassicas originally stated.The original
(1843, Beitr. 6, Pl. 6, Fig. 4). The form called
d e s c r i p t i obny D e s l o n g c h a m p( 1s 8 3 5 ,p . 7 4 , P l . 3 , F i g .
"Tetrdopsispirformis" Mrinst. (ibid. p. 58, Pl. 6, Fig.
1-3) dealswtth Tettdopsisbunellii and T. caumontii (p.
3) is doubtless identical with it, confirmed by an
76, PL.3, Fig. 4-5). The identity of the secondspecies
examination of the original specimensin Munich.
with the first was establishedby d'Orbigny (18a1, p.
Hence also Loligo pyriformis d'Orbigny (1845 [1855],
19). Collection sites were: Courcy and Amaye-sur-
p . 3 3 6 , a n d 1 8 4 6 ,P a l .U n i v . ,P l . 1 2 ,P a l .E t r . ,P l . 1 0 ) .
Orne (Calvados,France).Under the name L bunellii
This authorconsidersthe fonn in questionto be closely
Desl. are also the speciesmentionedby Fdrussacand
relatedto Loligo brevisBlv. from the Brazilian coast.
d'Orbigny 1835 (Pl. 1), d'Orbigny 1842 (Pal. fr. jur.,
Belotettthisacuta Minst is listed as B. subcostata b y d ' O r b i g n y( 1 8 4 5 :P l . 2 2 , F i g . 1 - 1 8 4 6 :P a l .U n i v . ,
P l . 1 , p . 3 8 ) , i b i d . 1 8 4 5& 1 8 5 5( p . 3 6 0 ,P l . 2 0 ) , 1 8 4 6 ( P l . 1 3 , P a l . U n i v . ) , C h e n u 1 8 5 9 ,p . 3 8 , F i g . 1 0 0 ; Keferstein1866,p. 1442,Pl.130,Fig. 1-2.
P l . 1 6 ,F i g . 3 , P a l .E t r . ,P l . 1 3 ,F i g . 3 ) . T h e s p e c i m e n is drawn like other figuresproducedby Miinster - using
A close inspection of the figures (in d'Orbigny
a mirror for symmetry, which explains the difference
1845)revealsthat the rather free reconstruction(Figs I
from my Figure 54. Occur:rence: Boll, uppermostLias
and 2) shows a shell of unique characteras far as the
e fl-ower Toarcian].Specimensin Munich and Berlin
outline and the growth lines are concemed.There is no
collections.
distinction between lateral plates and conus vanel nor
This shell form (p. 148) is closely relatedto the
of lateral arcuatelines in the latter; insteadthere is a
previous ones, but it could indeed be taken for a
very regular, leaf-like outline. The more realistic
separategenus. Characteristicfeatures in general are
depiction in figure 3 showsthe typical characterof the
the delicatetexture (which looks like a compressedleaf
beloteuthidsin that a bend in the growth lines slightly
in calcareousshales),the enhancementof the striation
demarcatesthe conus vane. However, this observation
radiating from the posterior pole, and the preservation
was rather tentative. In the Stuttgart natural history
of the ink sac. But it is the outline that provides the
collections,I found a shell with the following note: "In
most characteristicfeature: the anterior end is drawn
bituminous limestone with Ammonites.fimbricatus, Lias e fl-ower Toarcian] Courcy (Calvados)". This
out into a sharp point, much ltke Loligo (Fig. 58), the transition to the broad part of the pro-ostracum
specimenwas probablyexaminedby Quenstedt(1849,
thereforeis marked by a curved line, the lateral bands
89
JLT
Sy
Mesotcuthoidearesernblingrecenttypes. [Fig. 5a & 55] a. Beloteuthis acutoM:'i;nst.1843,reconstructcdfrom the original specimenin Munich (researchcollections).(Bcitr. VI. Pl. 6, Fig. 4). (From thc Upper Lias e, nearBoll). Fig.51.
b. B. libanrttic,/n. sp. (MS: the original specimenin Stuttgartis labelled "Geot. libanotica O. Fraas, Sahel-Alma"); posterior parl reconstructed. Cretaceousof Syria. Both I a nat. size. lb. ink sac.ao. lateralline. la. lateralasymptote.Between the two lie the "laterai arcuatc1ines". Fig.55.
Polaeololigo oblonga (Wagner 1848).reconstmctedfrom the original specimenin Munich (public collections)(Wagner, Pl. 24. Fig. 2). A shell fotm rather similar to beloteuthids;its outline could also be taken as typical for recenrmetateuthoids. r/5 nat. size.- Lithographic limestonesof Daiting. The dotted lines in the anteriorpart mark the approximateoutline of the pro-ostracumof related forms: I. Bel. acttta,IL B. bollenis. III. B. .subc'ostata, IV. Belentnites(or Trachvteuthis),withcorrespondingpositionsof the s t e l l a t e g a n g l i a ( G a/ 1n d G 1 2 \ . R i .k e e l o f t h e m e d i a n p l a t e ( r a c h i s ) R ; a . l a t e r a l l i m i t o f r h e m e d i a n p l a t e ( m e d i a n a s y m p t o tSer)i ;. . longitudinallystriatedzone ofthe lateralplate (Sp);Aa. lateralasymptotc;Ct. conusvanel Co. conus.(cf. Fig. 5g).
show very steepliee margins and growth lines (ia-aa).
be securelyincluded in the genusBeloteuthis,but I am not willing to create a new genus without having acquireda deeperinsight into the natural relationships
Problematic forms.
of this form:
The work of Konig (1825)was not availableto me, so I am not in a position to esrablishthe relationshipsof
Beloteuthis libanctica n. sp. Here belongs one shell from the Lower Cretaceousof
Loligo af/inis andlongaevus(Pl. 17, Figs 202 and 203).
Syria, now housedin the natural history collectionsat
The same is true of Ptiloteuthis.foliata (p. 150) which may belong here. Likewise the following form cannot
l i h a n o t i c a O . F r a a s , S a h e l - A l m a "( F i g . 5 4 b ) . T h e
Stuttgaft,with the following label: (147) "Geoteuthis
90 figure demonstratesthe lack of any relationshipto
Daiting, thus providing an exemplary case of shell
Geoteuthls, not even to the form placed in
morphology. One gets an impressionthat this shell
Trachvtettthis(p. 134); it is a beloteuthidsimilar to the
resulted from lengthening of the free rachis of
previousspecies,or at least a closely relatedtype. A
Belotetrthis actrta. The median keel (119) appears
differencefrom,B. acuta ts the ratherrobusttextureand
somewhatbroadenedbut is still accompaniedby the
the strong keel at a small size. The growth lines are
same zone of longitudinal striation. The lateral plates
indistinct, and the free margin lacks differentiations
show the samecurvedmarginsand are connectedto the
reminiscent of lateral strips or a conus vane. Perhaps
conus vane in the same way, with a sharp marginal
this incompletelypreservedshell belongsto a special
indentationand a distinct lateral asymptote(Aa). The
type. In any case,the short anterior tip of the pro-
spoon-shapedconusmust be reconstructedin a similar
ostracumplaces it closer to Beloteuthis acuta than to
fashion,basedon the observeddetails, i.e. as a spoon-
Palaeololigo.
shapedstructure.In the fossils it is destroyeddue to
However, transitions between this and the
flattening of the shells in the shales,but its curvature
following family must be consideredlikely, since we
can be reconstructed,thus leading to the present
are dealingherewith the evolutionaryline of the recent
representatlon.
teuthoids.Thus boundary definitions are arbitrary.
There is one remarkable difference: the lateral arcuatelines are not clearly demarcatedbut are (again) absorbedin the conusvane.
N. The family PalaeololiginidaeNaef 192I .
The rresoteuthoidmetamorphosis(p. 136) here seems to reach its full development, from which
In this family I consider mesoteuthoidswith shells
numerousother variantswere subsequentlyderived, as
similar to Palaeololigooblonga(Fig. 55).
shown by the metateuthoids. The anatomical
Diagnosis: Palaeololiginidsare moderatelyslender
significanceof the type is illustratedby the correlation
m e s o t e u t h o i d sw i t h L o l i g o - l i k e s h e l l s w h i c h e n d
between the rachis and the stellate ganglia (Fig. 55.1,
anteriorly in an elongate,stalk-like free rachis and
the position of which is clearly different in belemnoids
posteriorly have a leaf-shapedvane (conus vane plus
Glr and in metateuthoids.The muscular mantle of
lateralplates)with a spoon-likeconus.
coursefollows the recedingshell margin; so here we
The most striking featureof this shell is its marked
again see the strengtheningand concentrationof the
similarity to the gladius of recentsquids(Fig. 58). This
neuromuscular,active systems at the expense of the
similarity is so pronouncedthat one can ask whether
passiveshell apparatus.
we are looking at a closerelative of the recentgroup.A
The shell of Palaeololigo oblonga is about as
closer inspection, however, reveals the undeniable
delicateas that of Beloteuthis acuta. Its geological
relationship to the beloteuthids, especially an
occurrenceis limited to Malm s [E] of the Solnhofen Formation fl-orver Tithonian]. Its normal size is about
associationwith Beloteuthisacuta. (Seefurther below, p . 1 4 8a n d 1 5 4 ) .
15 cm. But smaller (younger) specimenshave been found.
The genusPalaeololigoNaef 1921.(System,p. 535).
There are several problematic forms from the Palaeololigo context: an apparently related form is
Here belongs"Teuthopsis"oblonga Wagner 1860 (p.
T u s o t e t r t h iLso g a n ( 1 8 9 8 ,p . 4 9 7 ,P l . I 1 0 , F i g . 1 ) , f o r
50, Pl. 24, Fig. 2) and T. princeps (ibid., Fig. 2). My
which I have only indicationsby v. Btilow (1920, p.
examinationof the specimensdescribedby Wagner
262). According to this author, the shell is oval in
convinced me that they representa single species,
outline and has a median keel and a long free rachis.
which must cany the first namecited (cf. p. 130).
The original descriptionwas not availableto me.
(r48)
Very questionable: Palaeololigo oblonga Wagner (1860).
Figure 55 shows a careful reconstructionof the shell based on specimensfrom Solnhofen,Eichstiitt and
The genusPhylloteuthisMeek and Hayden 1860.
91
o
k
p
h V
t .
/
a ..
: i i , . '-
h
:,::t
tr.*
r<
t
Fig. 56. - Celaenoconica (a, d) andscutellaris (b, c). ,r. Dorsalview, drawn from the original specimenin Munich (wagncr 1860,pl. 24,Fig.4-5). tri. Crosssectionofrachis.l/1nat.size. l' Shell with animal, compressed"seen in lateral view. From the lithographic limestonesof Blumenbcrg near Eichstiitt. Original :lrecimeninthecollectionsoftheLyceumof EichstZitt. ar.armbase; frp.head;rr'.funnel;lD.inksac; 77.stomach(thespiralstriation is very distinct, also seenin anotherspecimen;one tends to think of a swallowed ammonite,since a correspondingstructurein the "spiral caecum"is unlikely to be so u,'cl1preserved);rt. fuirnel retractor;rnu. manrelmusculature;ra. rachis.l/2nat. size (samefor c, and d). r'. Shell in lateralview (Munich, scientificcollections). r/. Profile of a specimenpreseruedin three-dimensional form (Munich, idem) t' Acanthotetrthisproblematica (cf. p. 183) liom the lithographic limestonesof Daiting. The phragmoconeis compressedstarting tiom the apex ('t,.1',z); below it lies the ink sac (t); thc pro-ostracum(p) showsthe centralrib (lll); one can recognizethe head with eyes(a) and mandibles(t). and armswith hooks (i) (Fig. 68a).
Here belongs PhylloteuthlsMeek and Hayden 1860,
Here belongs Pt. .foliata Gabb 1869, a shell not yet
1 8 6 4 ,d e t a i l e dd e s c r i p t i o no n l y 1 8 7 6 ,p . 5 0 5 , P l . 3 3 , F i g .3 c f . Z i t t e l 1 8 8 5 p , . 5 1 9a n dF i s c h e r1 8 8 7 p , . 353.
figured.I quote from Fischer(1887,p. 354): "Gladius sub-oval without a median keel; surface showing
(150) This is an impression in a Cretaceousrock w h i c h i s r e m i n i s c e n to f B e l o t e u t l l s , t e n t a t i v e l y
region,partly from the centreof the blade." Neocomian
reconstructedaccordingto the aspectof a Palaeololigo.
of Califomia.(cf.Zittel1885,vol.II, p. 519).
oblique striations,partly radiating from the anterior
It is a very dubious fossil lacking any definite teuthoid feature.It is certainlyno closerelative of Palaeololigo. Indeed, the part interpretedas the vane shows a fine striationon either side,which is perfectly straightlike a
O. The family CelaenidaeNaef 1921. (System,p. 535).
simple hatching. The strips either side join in the middle in a weakly protracted angle.
E,qually
The characterof this new family is particularly unique
problematicis:
and its seprarationtherefore perfectly justified. One could even doubt its connection with other
The genusPtiloteuthisGabb 1869.
mesoteuthoids.But considering that the order should emphasizeconnectionsrather than differences.we
92 Fig. 2). Both namesare acceptedby Quenstedt(1849, p. 522).Wagner (1860,p. 32) correctlyusesonly the Itrst one.This species(Fig. 56b, c) is more robust than the following one and also differs strongly in outline. The shells of this speciesin generalare embeddedlaterally and thus offer a profile view as drawn in Fig. 56b-d. We can distinguisha strong rachis next to which (like in the handle of a spade)lie the lateral plates,forming an obtuseangle (about 120-130'). The margin, with two prominent angles, grades into the shield-like, bluntly pointed conus vane. The conus is blunt and sunoundedby the part formed by the lateral plates, so that a ridge appearson either side of the laterul (152) asymptotes. The rachis crosses this delta-shaped
\\ Fig. 57.
Celaenoteuthis incerta n. g., n. sp., from the
lithographic limestones,drawn fiom a specimen in the collectionsof the Berlin Natural History Museum (Fischer collection), added infbrmation from other specimens (Munich, Eichstiitt).r/2 nat. size. a. dorsal view; r. lateral
anterior parl like the ridge of a rather flat roof. I have seenwell preservedimpressionsof the soft parls of this species(Fig. 56b). They show, of course, an extremely plump animal, which was probably laterally compressed,with a thick head and mantle sac. The musculatureof the arm bases,the mantle and the funnel has left distinct traces. The ink sac also is
asymptote;b. profile (longitudinal section),c. cross section,
preserved.From the lateral angles strong muscles
slightly behindthe positionmarked,t,anteriorto the apex.
extendto the funnel, so they are easily identified as the funnel retractors.In the position of the stomach one finds a peculiar radial-spiralstructure,which could be
refrain from a more drastic separation.(Cephalopoda,
d u e t o a c o m p r e s s e d ,r i b b e d a m m o n i t e l y i n g
vol. I, p. 47 and147).
underneath.One might also think of swallowedprey or
D i a g n o s i s : M e s o t e u t h o i d e aw i t h a g l a d i u s characterizedby a long rachis, in which the conus did
of a spiral caecum. The last interpretation would be rather daring given the delicate nature of the spirally
not occupy the end of the mantle sac but was shiftedto
arrangedstructures(glandularfolds). There is no trace
the dorsal side (to a position comparableto limpets)
of arm hooks, either in this or in similar specimens. (But seeFig. 56e,Acanthoteuthis problematica).
and ventrally contributedto the lengtheningofthe vane in a posterior direction. - The following genus has been known for a long time; as long as it appeared isolated it was particularly problematic. I recently found another, very distinct shell type that has to be
b) Celuenoconica Wagner 1860. For a comparison see the original description by
consideredin this context.
Wagner (1860, p. 35, Pl. 24, Fig. 4) and Keferstein (1866, p. 1447, Pl. 130, Fig. 3-4) and my Acanthoteuthisproblematica. - Once Wagner had
The genusCelaenoMtinst.5l1842.
rightly united rhe rypes of Mtinster (cf. p. 15l), he referredto this new speciesas "Celaeno conicd', based
Here belong shells like Celaeno scutellaris Mrinst.
on rather heterogeneousmaterial. After removing our
(Beitr.5, Pl. I, Fig. l, p. 95-96).
Acanthoteuthisproblematica a form of celaenid shell remains (Fig. 56a, b) that is rather different from C.
(ts1)
scutellaris. The overall aspect is markedly more a) Celaenoscutellaris Mtinst. 1842.
In addition to the type cited (loc. cit.) C. arcuata (Ibid.
delicate, the shape is broader, and the embedding in general is dorso-ventral,permitting three-dimensional
93 -fr'servationwith only moderate compression.The :.rchisproper is narrow and delicate, adorned with
probably closer to the general norm fof the Metateuthoidea,presumably]than the other forms. The
.:rcuategrowth lines. The lateral plates are subdivided
next closestrelative seemsto be C. conica, whereasC.
::troseveralpartsby longitudinallines that radiatefrom
scutellaris representsan extreme state.
:rc conus:on either side of the rachis a narrow strip
The subdivisionof the pro-ostracumindeed is very
-'rtcnds far forwards, whereas the rest is shorter,
similar Io C. conica: the lateral plates are subdivided
-'rtendingfrom the rachis at a very obtuseangle.If we
into severalsections,althoughthe part lying next to the
:nterpretthe line firarkedx as the lateral asymptote,the
naffow rachis does not reach further anteriorly than the
.'onnectionwith the broadly oval conusvane is formed
more lateral ones.We again considerthe line markedx
rv an indentationor a depressedradial groove, in :rarkedcontrastto (153) the previousspecies.(But we
as the lateral asymptote,next to which lies a distinct "lateral arc zone". The blunt lateral angle supposedly
:ray have assumeda false homology in that the line y
belongs to the conus vane. Rather inconspicuous
:nieht correspondto a radial line not drawn here. If so,
grooves radiating from the conus underline this
lhe interveningsectionwould representlateral strips ; 'en.suFig. 53r) seeminglyintegratedinto the conus
subdivision and demarcatethe part of the spoon lying behind the angle just mentioned.This spoon-shaped
r ane in C. conica, in the lateralplate in C. scutellaris).
part is particularly delicate and is always torn, either
.\t any rate, the two speciesare markedly different. If
due to compressionor even before burial. The anterior
creationof a new genus should becomenecessary,I
parts are more robust and adornedwith wavy, widely
ri'ould suggestthe name Listroteuthis(from )"totpov:
spaced growth lines which appear knobby, allowing one to recognizeeven small fragments.- Localities
ladle).
were Eichstrittand Solnhofen. The genusCeluenoteuthrs nov. gen. \Vith one new species:
P. The recent MetateuthoideaNaef 1921 (System,p. 535) and their relationship to the fossil Teuthoidea.
Celaenoteuthisincertu n. sp. Figure 57 showsa shell clearly reminiscentof Celaeno;
We have seen that forms strongly reminiscent of the
some of the fragments I found in the Munich collectionsare indeedso labelled(a1I . I) (b2).These are peculiar indeterminate remains, which are
recent relatives of the genusLoligo are already found in the Mesoteuthoidea(p. 1a8). One might even be tempted to use this similarity to suppressor move
recognizablein a generalway without allowing one to
traditional systematicboundaries.But then the genus
precisely reconstruct the whole. I found a fine
Palaeololigo would be united with the Metateuthoidea
specimenin the collectionsof the Lyceum at Eichstritt,
and would drag the beloteuthidsand trachyteuthids
labelledas "Ostracoteuthis superba" Zttt. (?)". The
along with it, thus obscuring the homogeneousaspect
best specimen is housed in the collections of the
of the enlargedgroup; but there are also other reasons
Natural History Museum in Berlin; it is from the collectionof V. Fischer(part and counterparl).It shows
for keeping the recent and fossil groups apart, in spite
the shell compressedbut still in a three-dimensional
situation is indeed very different from that in the
of their obvious relationship: in the teuthoids the
state; I could combine it with sketches of other
sepioid type, which indeed is the result of a
specimensto give a completereconstruction.
rearrangement and new creation.The mere reductionof
From the lateral aspect(b) one can understandthe
the conusand its phasesare not such conspicuoussigns
derivation of the celaenid type from common
of close relationshipas are the general featuresof the
mesoteuthoids.The muscularmantle appearsextended
sepioid shell. As to the details,one cannot deny the possibility that the occuffenceof a truly conical conus and ofa (156) distincl rostrum in recentrnetateuthoids
in the posterior direction, along the margin of the conus.This openedthe possibilityof moving the whole conusto the dorsal side, while the margin of the conus permitted further growth of the vane (154) tn a
makes their derivation from older (prototeuthoid)
posterior direction. The present celaenid form is
therefore,derivation from mesoteuthoidsis not an
forms at least conceivable;in phylogenetic terms,
94
Bd
3 ,f*t
N ij
!
'Kq
,
.r
!'l 'l:lt' ;
)l \,,'lif (
,r,.i.;_
'.
\::i-/
)
Fig. 58. The morphologyof Loligo vulgaris Lam. (Recent). 1/2 nat. size,drawn from life. Il2nat. size,also for the figures c, d, e; b. Frontally openedmantle sac with organsremoved, revealing the insertion ofthe shell.
a. Ventral view of a maturefemalefrom the Bay of Naples. figure./r/1nat.size.
c. The anteriorpart ofthe body and the head-foot,taken from the preparationshown in b, in dorsalview. The gills are slightly spread out; normally they would be alignedwith the marks 1-J, and the branchialbands(KD) would lie againstthe inside of the mantle.The shell epitheliumand (in zoner') thc primary mantleare visible; the mantlecavity (-x)is alsorecognizablebelow the mantle. d. The preparationof c in lateralview; the anteriorpart of the primary mantle (Ra) bearingthe nuchal attachmentis cut medially; for
(r\ the rest the whole extentofthe epithelialsurfaceofthe shell is againrecognizable. c'.Crossscctionsthroughthe gladiusof a somewhatlarger specimen.f . in thc areaof the nuchal attachment(Kg); 2. betweenzones2 and J; J. at the broadestpart ofthc vane. l. Yonng loliginid (AlloteuthismediaL.). Po. orbital pore; Pu. pupil;1r. iris lobe; Ro. olfactory organ;H/. nuchal cross fold;11o. knob situatedbesi
indisputablescientific assumption.lt is even more questionablewhether theparticular featuresof the soft body which are characteristicof recent teuthoids are
intermediatestages.(Seethe Oegopsidain particularin the forthcoming parts of the monographCephalopoda,
a l s o r e p r e s e n t a t i v eo f f o s s i l m e s o t e u t h o i d s t; h u s
vol. I, chapter9). More important here are the primary juvenile stages(cf. Figs 60 and 61) which must be
abandoningthe separationwould also jeopardizethe
consideredtypical.
currentfeaturesofrecent squids. Diagnosis: The Metateuthoideacomprisethe recent
The typical conus, which is never absent from advancedembryonicand earlyjuvenile stages(Fig. 60)
teuthoids in which the gladius shows a long "free
can also undergo a secondary
rachis", the posterior continuationof which in general
inexperiencedobserver will not be able to find it by
is accompaniedby a leaf-shapedvane, which in turn is lrore or lessdistinctly separatedfrom the conusvane,-
dissection,e.g. in Lctligo vulgaris and close allies. lt
in which the posteriorend of the gladius,at least in earlyjuvenile stages,bearsa conical to spoon-shaped
this state.Given the small size of a juvenile, the conus
- in which the buccal pouchesare very deep conus52,
identification is therefore of great importance.ln the
and together form a slit-like spacearound the buccal
Oegopsida(Fig. 61a) it is in generalmost completely
mass,the six compartmentsof which are separatedby
developed at early post-embryonic stages; it thus
thin membranesbut communicatewith one anotherin their deepestparts5r,- which are"non-stop sw*irnmers"
provides an important factor in the context of our
leadinga purely nektonic life,
the muscularmantle (cf. p. 22). But it is not necessarily
which lay their eggsin
redttction. An
often is very small and delicateand usually remainsin is a minute elementof the adult gladius.lts ontogenetic
conceptionof correlationsbetweena conical shell and
(157) capsulescontaining at least four, generally many
inhibited in its furlher growth to finally disappearfrom
more eggs.(In living sepioidssingle eggsare spawned:
the overall aspect of the gladius. In many casesit grows slowly to the adult stage and then forms a
F i g .3 8 c ) . (158,)Figure 59 illustratesthe special morphology
considerablepart of the shell (Fig. 59) as is typical for
and topography of the gladius of metateuthoidsand
many families. In such casesit conservedits primary
also provides a basis lbr reconstructingmesoteuthoids.
relation to the muscular mantle, i.e. its free margin
A typical featureof developmentin recentforms is the processby which the muscularmantle progressively
nrantle (Figs 10, 39, 42 and 58). (See Cephalopoda,
shifts its insertion, in general during post-embryonic
vol. I Pl. 4). In some large metateuthoids,the small
development,to the outside of the shell, so that the
conus can indeed be recognizedby palpation of the
shell can finally become entirely envelopedby the muscularmantle (Fig. 7e). This latter stageis always a
[posterior]end of the mantle,if it is not visible through the surfacetissuesas in delicatejuvenile forms. When
secondarystate, and the developmentleading to it appears(in different species)to be halted at different
a distinct rostrum is lacking, the conus is coveredby a fleshy terminalend.
offers a permanent insertion site for the muscular
96
F
H a B 3
gt EI
t'1 lll
WN
Wt Fig. 59. Morphology and topographyof the gladiusin recentMetateuthoidea. a. Ideal prototype.Insertion of the gladius into the muscular mantle (Mm) the insertion of which (Ma) continuesal1round the free margin (Cr). Relationshipof the vane (Fa) to the fins (F/) and fin cartilages(Fn); Fh. membraneouspart, connectedto the rostmm and ccnus;O/. ear lobe; Rp. rachispart ofthe vane;R/r. free rachis;De. dorsalprojectionofthe mantlemargin. a1.Ventral view of the anteriorend of the mantle. d2.Ventral view of the posteriorend of the mantle.Nfr. nuchal attachment;Ve.Yentral angle;Ta.funnel notch; Fr. free margin of the posteriorpart of the gladius;Co. conus;Rs. rostrum,supportingthe terminalpoint (.!p) of the mantlesac. b. Young individual of Gonotus.fabrlcll(Licht.) from Bergen.in dorsalview. 2 1 nat. size.The muscularparls (muscularmantle,fins) arejoined abovethe gladius;only the rachisis recognizablethroughthe skin. b1.Posteriorend in ventralview, with the conusvisible throughthe skin. c. Gladiusof the samefotm, ventral view. 2t l2 nat. size. -l. free rachis;2 . lateralribs of rachis;J. median asymptote;4. lateralplate; 5. stiffeningrib oflatcral p\ate;6. margin ofconus vane dying out anteriorly;7. conusvane;B. conus. cr. Conusof the abovegladiuswith phragmocone-likefilling, in lateralview, after Steenstrup(1881,Pl. l, Fig. 7).2/1nat. size. d. Gladiu-iof a young individual of Sthenoteuthisbartrami (Les.) with totally degeneratelateralplates,in ventral view (lateralview of the endof thegladiusnextto main figure).| 2 nat.size. lr., 1r. doublereinforcing ribs of rachis (Rl). In the posteriorpar1,the vestigial lateral plate forms a third elementaddedto this rib. Sl. stem;Fa. conusvane; Co. conus.
97
Fig. 60. - An embryo of Loligo t,ulgaris in semi-schematic(median) sagittal section.'u/r ndt. size. Inside the shell sac (.tr) one can recognizethc delicategladius with a distinct conus.Inside it a specialconcentrationof cells (Sl) indicatesthe rudiment of a siphuncle.which soon degencrates. Ra. rachis of the gladius. enclosedin a namow. thick-walled part of the shell sac.11o. Hoyle's organ (longitudinalbranch);lo. anterior aorta.Md. dorsal mantle cavity n'ith nushal attachment(N/r): Oe. oesophagus;Grl. poison uland; Cg. cerebralganglion:Rd. radular pouch; Ob. upper buccal ganglion; U1l. lower buccal ganglion; Ofr. upper mandiblc; Uk. lorver mandible; 11.inner lip; ,4/. outer lip; Mu. primary mouth: Do. yolk; Sr. subradularorgan: Pg. pedal ganglion; Sn. (vcnous) sinus; lg. visceralganglion;Sl. statocyst;Vc. vena cava; Td. ink glandt 81. caecum:Gg. ganglion gastricum;L1a.stomach:C6r, Cii), Co1.parts of the coelom; Go. gonad; Gr. genital veinl F1:.heart;Nl. kidney; rUs.mantle septum;Mnr. muscularmantle: Mr'. mantle cavity; Af. anus:Dr. funnel gland: Tfr.funnel valve; fi'. funnel tube. still closed by a membrane.
In some families one finds problematicremains of a
DiploconidaetAmblvbelu.s| andFig. 71v).
f o r m e rp h r a g m o c o n e .( S e ee . g .p . 1 0 4 a n d t h e ( 1 5 9 )
Most metateuthoidsshow a strong (161) secondary
explanationsto Figs 78 and 19). A deposit of shell lrateriai in the conus is very comlnon and can indeed
lengtheningof the./ins, which at post-embryonicstages grow forward along the sidesof the mantle.During this
be comparedto a phragmocone.A detailed study has
processthey necessarilygrow beyond the area of the
not yet beenmade.However,Steenstrup (1881)already
conusand conusvane and thus reachthe surfaceofthe
fbund true septain Gonatus,an observationconfirmed
muscularmantle.Even if the muscularmantle doesnot
and supplemented by Hoyle (1889,Figs 39c and 59c1).
envelopethe shell, the fin baseand its ctrticular pouch
The material depositedin the conus doesnot appearto
(Fig. 59a) comesto lie on a new supportwhich does
be pure shell rnaterial, however; part of it is
not lirnit further extension until the anterior mantle
c'artilaginous tissue, as is often observedin the shell
rnargin is reached. Apparently a similar process
complex.Moreover, there seemsto be no indication of
occurredin the sepiids(Fig. 31) and in the convergent
a siphuncle(cf. Fig. 60); a strict comparisonwith the
genusTrachvteuthis(p. 139). Individual speciesof
belemnoid phragmocone thr.rsappears impossible.
rnetateuthoidsshow diverse stagesbetween the initial
Nevertheless,the similarity of Steenstrup'sfigure with
situation and the potential final state.lncidentally, the
a problematic fossil describedby Krorr (Suppl, Pl. 4f,
formation of an ear lobe (Fig. 59a) at the anterior end
Fig.2) is strikingB ; l a i n v i l l e( 1 8 2 7 ,P 1 . 3 ,F i g . 1 4 ) ,a n d
of the fin permits lengtheningbeyond the anterior
d ' O r b i g n y ( 1 8 4 6 ,P a l . U n i v . , P l . 7 7, F i g . 1 0 -I 1 , P a l . E t r . , P l . 3 7 , F i g . 1 0 - 1 1 )p i c k e di t u p a n d n a m e di t
mantlemargin (Sepia).
Belemnitesobtusus.Perhapsit is merely a composite
Such a lengthening of the fins is related to an e n h a n c e m e n ta n d e x t e n s i o n o f t h e i r . f u n c t i o n a l
picture that could be misleading.But it could also have
significance. Whereas the primary short, lateral-
a real basis. At any rate, that is the way one can
terminal extensions(Figs 42, 43 and 6l) can function
imagine the transition from a true (160) phragmocone
like a rudder and elevator,avoiding rotation of the
with degenerationof the rostrum to a sirnple filling of
body aroundits longitudinalaxis (p. 35), elongatedfins
t h e c o n u s , a s r e p r e s e n t e di n F i g . 5 9 x . ( c f . t h e
become active locomotor organs by using an
98 Fig.61.
Young teuthoidsfrom the plankton of
the Bay of Naples(oegopsid"lar-vae"). a. Pyroteuthis margaritifbra.Ell nat. size. Ventral view of preservedanimal. b/. blue light organ:g/. greenlight orgar.. at. The same,from sketchesmade from the live animal. a2. Dorsal view of a. ld. lld pore. Note (visible through the skin) the dark gladiuswith a spoonshaped conus, the insertion of the muscular
o.
mantle on the free edge of the shell, and the insertionofthc fins on the conusvane. D. Young individual of Onvchoteuthis banksi. In,
" 1 r 1 ? t s. i z e .V e n t r a lv i e w . T h e c o n u s( C o ) i s markedly degenerateand shows a distinct.
cl
pointed rostrrm. Ift. funnel attachment.Note the four, sti11 undeveloped ventral arm rudiments. b1.Dorsal view of the same. h2. Lateral view. The mantle is shrunken,thus exposing the posterior parts of the funnel apparatus, which are normally hidden. Zl. funnel pouches;Nft. nuchal attachment. l0/1 c. The same speciesJsomewhatolder stage. nat. slze. c1. The isolated posterior end, at greater magnification, in lateral view and (optical) sagittal section. -1. margin of conus vane; 2. cavity of conus; 3. pro-ostracum in sagittal section;/. rostrum. 8/1 d. The same species,further developed. nat. size. The ventral part of the muscularmantle is cut away.
0.2
lt. ventro-lateralann; la. ventral ann at early
b.
b.
growth stage; Te. Lentacnlararm; Vc. vena cava;
Il. funnel attachment;fi'. funnel retractor;Ib. ink sac;Kb. branchialband; Nl. renalpore; tr/s.venousappendages ofthe right branch of the vena cava;Kh. branchialheart; Vl. vena pallialis lateraiis;Km. gilllamellae; Rc. insertionof gill retractoron the rnantle;lo. posterioraofta,n. line to which the vane of the gladiuswould typically extend(and to which it indeedextendsin relatedspecies);I/p. venapallia1isposterior;Ap. arteriapallialisposterior;Co. conus;Rs. rostrum;Ms. mantleseptum.
Ltndltlatingmovementof the fin musculature.Of course intermediate stages and transitions also exist in
Jurassic.They first occur in Lias e fl-ower Toarcian] with numerousspeciesand severalfamilies (p. 111,
functional terms (cf. Cephalopoda,vol. I, chapters6-
122, 125, 132 and 141), mostly representingancient
31).
types often with great similarity to belemnites;among thesetypes some alreadytend to point in the direction of Loligo.
Q. Review of the evolution of the Teuthoidea.
(162) A chronologically sequential gradation cannot be established,however. In Malm e fl-ower
Squid-like shells are still absent from the Triassic 54,
Toarcianlmore definitely Loligo-ltke shells (p. 148)
but they are present in diverse forms from the early
already occur, similar to those (Palaeololiginidae)
99 found in the Upper Cretaceous(Senonian).In Middle
evolution was an increase in li/b intenslf acquired
Jurassic and Tertiary strata conditions of marine
through liberationfrom stereotypesand renunciationof
sedirnentationwere apparentlynot appropriatefor the preservatiorr of similar types; in more generalterms,
passivesecurity;this point ofview of courseappliesto any "ascending series". Elsewhere we will try to
only a small.fraction of exttnct speciesis representedin
analysesuch a progressionby successivegradesand to
the fossil record. The known forms thus can only be
draw some biological conclusions.Here the observed
consideredas landmarlrs, never as evidencefor a
facts can only be given in their systematiccontext, in
continuou,s h istor.v. Even so are they of great
the light ofgeneral insights.
signifrcance,as they provide somemajor links between the living decapods(squids) and the predorninant Mesozoicforms (belemnoids).Let us try to visualize this link through a series of real species,without claiming a truly phylogeneticorigin: 1. ln the Upper Triassic we find aulacoceratids with a reduced phragmocone:Calliconites dieneri (q. v.). 2. In the Lias e fl-ower Toarcian]we find prototeuthoidssuch as Geoteuthissimplex (p. 123). 3. Apparently relared to theseare the contemporaryBeloteuthi,ssubcostataand 4. B. bollensls (p. 143), initiating a series of more
(164) Part IV: The Belemnoideaor belemnitelike dibranchiates. Content.s'. A. Preliminary remarksand Diagnosis(p. 165). B. On the typical structureof belemnoids(p. 165). a) Structure of the shcl1(p. 167). b) Materials for reconstructionof the
closely related speciesthat continues5. through the
soft body (p. 176).The genusAcanthoteuthis(p. 177). The genusBelemnoteltthis(p.185). c) Generalaspectsofbrachial
d o r . r b t l e s sy o u n g e r B . a c u t a ( p .
146) and 6.
armament(p. 187). d) On the position of the shell inside the
Palaeololigooblonga (p. 1a8). The strong similarity of
soft body of the animal (p. 189).C. On the lLnction of typical
the last form to recent shells is striking, as has been
belemnoidshellsand the life style oftheir bearers(p. 191). D . T h e l a m i l y B e l e m n i t i d a e( p . 1 9 3 ) . E . T h e t a m i l y
emphasized above(p. 156). lf we ask whether this metamorphosiscan be consideredfrom a generalpoint of view, we can give a positive answer based on what we have seen for sepioidevolution (p. 95-96): the shell reduction, which
Phragmoteuthidae (p. 260). F. The family Aulacoccratidae(p. 262). G. The family Xiphoteuthidae(p. 214\. H. The farnily Belemnoteuthidae(p. 276). I. The tamily Diploconidae (p. 278). K. The family Vasseuriidae(p. 280). L. Review (p. 281).
can be followed through the palaeontologicalsequence
I i
i
up to the divergent fonns within recent groups, signifies more than just a direction-lessdisintegration
A. Preliminary remarks.
of an older form, it indeed also shows an lncrease (made possible by surplus energy) of the effective
There is a rich literatureon belemnoidcephalopods,so
range by means of an enlargementof the muscular
one might assumethat ideasaboutthis group havebeen
mantle. The animal progressively rids itself of a clumsy, demanding hydrostatic and protective
clarified. A closerlook howeverrevealsthat zoological observers lacked sufficient palaeontological
apparatusthat is essentially passive in nature, to
knowledge, and vice versa. In fact, neither the
ultimately rely, as an active individual, on the
comparableliving nor the fossil representativesof
voluntary use of its weaponsand tools in the struggle for existence.We here seea continuationof (163) the
sepioids and teuthoids have so far been studied in
changesthat had allowed tetrabranchiates (p. 24) to be
terms, to allow one to base the reconstructionof
transformed into dibranchiates. A
belemnoid remains on such knowledge. It must be
preliminary
sufficient depth, in systematic and morphological
condition of both augmentationswas the superior
emphasized therefore that fully
differentiation of the muscular, nervous and sensory tools, the unused potential of which opened new
reconstructions of belemnoid animals have not been producedto date. Even the most recent ones by O.
avenues.Old requirementshave thus becomeobsolete.
Abel (1916) lack a sufficientmethodologicaa l nd
One gets an impression that the "sense" of this
materialbasis,even though a considerabledegreeol
substantiated
100 insight into natural history is recognizable,with a
animals and thus leaves little doubt about the overall
serious attempt to penetratethe morphological and
picture. (166) As to the details, we will try to justify
ethologicalessence(165) of the whole group (p. 7).
our view.
The systematic-morphological tools, which I have acquired during the previous six years of work, are
a) Structure ofthe shell.
indeed indispensable,as are observationsof living
Complete belemnoid shells unfortunately have never
anirnals.Even the most careful worker will be misled
been found, so tedious studieshave been necessaryto
by illustrations and literature repofts if he is not fully
elucidate the structure of the shell. Although our
familiar with the field in question.There are countless
presentideas seem correct, at least in their general
biased and fal.se ideas about the rather casually treated
outlines, since they are based on a great diversity of
i n v e r t e b r a t e s , w h i c h a r e i n d e e d d a n g e r o u sf o r a
proven facts, they still need improvements and
comprehensivetreatmentlike the one attemptedby
additionsin someessentialparts.
Abel. A
The phragmocones(p. l5) have beenknown for a reconstructive palaeomorphology and
very long time, and they have correctly been compared
palaeoethologycannot be based on circumstantial
with the older nautiloids(orlhocera),sometimeshaving
evidencederived from overall similaritiesor
even
been confused and mixed up with them. This is
worse from the shapeof the posteriorend.A striking,
perfectly understandableconsideringthat they lack
but partly atypical similarity may occur together with
distinctive featuresand often occur separatedfrom the
very marked differences. I do not understand, for
parts of the shell which show more specihc characters.
exarnple,how one can assumethat the rostrum-bearing,
Their shape is variable as in orthocera.The cross
calcified shell of a belemnitecan occur in a delicate
sectioncan be circular or oval, and in the latter casethe
gelatinous,nekto-planktonicdecapodlike Chir othauma
smaller diameter may be orientateddorso-ventrallyor
(Abel 1916, p. 173), going so far as to add the
transversely,depending on the species.The apical
membraneousaccessoryfins and other details of this
angle also shows great variation: in the most slender
extravagant, absolutely atypical form
phragmocones it is about5o, in the stockiestones30o.
to
a
"reconstructed"animal (Figs 78 and 80). The stronger
The total length of
body of an Alloteutlrls (Fig. 74) may perhapsbe suitable for such a pulpose (Fig. 80a), likewise the
phragmoconesranges from I cm to about 40 cm, the t/,* diameter of the protoconch from to I mm. As a
body of a Sthenoteutlrls(Fig. 80c). But in principle,
generalrule the phragmoconesare very weakly curved,
this method of dealing with the problem is highly
the marginal siphunclelying againstthe concaveside,
controversial.(Seeour principlep. 7).
as in sepioids(part II, p. a5). The concavesurfaceis
known, well
preserved
Diagnosis: The Belemnoideaare fossil decapods
thus the ventral side, as in the sepioids;in most cases
with a well developed,straightphragmoconeat the end of the mantle sac,- which appearto have transformed
this can directly be derived from the shapeofthe shell,
the suckersinto hooks on some or all arms.- Probably
positionofthe pro-ostracum (Fig. 71).
and in the belemnoidsit can also be deducedfrom the
the earliestbelemnoidshad normal suckersthroughout
The phragmoconeis often preservedwith its typical
their lives; at the presenttime we know nothing about
envelope,the conotheca(p. 15). It showsthe normal
their brachial armament; only the hooks of younger
two layers, which do not apparentlydiffer from those
types are known.
of common orthocera;in most instances,however,they appearextremelydelicate.Moreover,there is no coarse outer sculpture:the ostracLtmis always rather smooth
B.
On
the typical structure of
belemnoid
dibranchiates.
and only shows very fine lines (grooves or ridges). Theseoccur in two groups.There are longitudinalones which convergetowards the apex of the cone (cf. Figs
Figure 62d illustrates our ideas about the typical
41, 45, 41, 5l), (168) and there are curved lines
structure of a belemnoid. Although this figure
showing a complicatedcoursein such a way that each
representsa morphologicalreconstruction,it is very
individual line runs all around the cone close to the
close to well preservedfossilsor impressionsof such
preceding and following lines. These are the grov,th
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iq \7 l
f
Fig. 63. Fossilsusedfor the reconstmctionofbelemnoid organisation.a-ft l2nat.size. a. Rostrumand phragmoconeof Belentnites" elongatus" after Quenstedt( I 849, Pl. 24, Fig. 3) fiom the Lias of Swabia. 6. Pro-ostracumfrom the Upper Lias of Alderton (Gloucestcr)after Crick (1894, Pl. 9), lower part reconstructed,to illustrate its origin on the phragrnocone.1. asymptote;2. centralrib; 3. f'eatherystriation: 4. Iateralplatel 5. annulus:6. anterior suture; 7. conus rim. c. Phragmocon e of Bel.paxillosus,in dorsalview; ,u.paraboliclines,.t. medianasymptote. r/. Split rostrum of "8. elongatusMiller" from the Lias 6 of Breitenbach(Wiirttemberg)(BavarianStateCollections)with the imprint of the sameparaboliclines in the alveolus. e. B. elongatuswith phragmoconeand ink sac on a slab of shalefrom the English Lias (aftcr Huxley 1864).The dotted arrow, when doubledin length,nould indicatethe position on the plate where a seriesof belemnitehooks is situated.At this point the anuscan be assumedto lie. Thus the hooks belong to a coprolite rather than to the arms of thc animal itself (rememberthat cephalopodsarc cannibalsl). /. Shell of "Aconthoteuthisspeciosa" from the Upper Jurassicof Solnhofen.Specimenfrom the Munich Museum (Bavarian State Collections),with complementaryinput liom other specimens.Such fossils also occur togetherwith soft parts (as in g). ,/. proostracum(medianplatc).2. median asymptote,S. annulus,4. anterior suturelinc of the last septum,5. posterior suture line (mural ridge), 6. idem to pcnultimateseptum.7 like 2 . g. Body of "Acanthoteuthisspecio,sa"from Eichstiitt(after Crick 1897,P1.1). a. arms. 6. mandible,lying (as is often observed)in a 'i2. group of calcareouscrystals,c. head./. liver (?), L ink sac.rr. muscularmantlc; at the left: a hook in naturalsize.othelwisenearly
=
103 Fig. 64. Typical affangementof the shcll and mantlein decapods. o. Shell location. The mantle is empty, the lateral pafis of the muscular mantle(Mm) and associatedstnrcturesincluding the shell being prcserved. Anterior to thc line :r the prirnary mantle is present,covering the shell and cxhibiting a spccialstructureacting as an adhesiveand sliding surface(G1): thc often cartilaginous "nuchal attachment". On the pro-ostracum some growth lines are shown as dotted lines. Np. nervus pallialis; ,St.stellate ganglion;Kb. branchialband; Vl. vcna pallialis lateralis;Fn. ftnnewe; Vp. foramen lbr vena pallialis posterior; lp.
foramen for arteria pallialis
posteriorto thc fin; Pl. phragmocone,exposed. b. Mantle locotion. Natural topographyafter removal of the ventral pafi of the muscularmantle.Ro. oli'actory organ;Th. funnel attachment;Zr. funnel pouch (the "funnel tube", which is the main part of the "funnel apparatus", lies in its centralpart); Vc. vena cava:Tr. funnel retractor;Erl. intestine;Go. gonoduct opening; Ra. musculus rectus abdominis; Ic. accessory nidamentalglands;Mp. renal pore; Nrl. nidamentalglands; Vl. venapallialis lateralis;Kv. branchial vein' Kh. branchialheart;Pd. pericardialglandl Co. coelomic pouch for Pd: Mt, M,. Remainsof primary mantle, on the inside of the shell; Am. arteriapallialis medialis; Vp. vena palliatis posterior;lp. arteriapallialis posterior:.tr/.fin.
Thesefiguresrepresenta conectedversionofthc samefigureson p. 124 in"Cephalopoda",vol. I; it now appearsthat the fins are to be placedon the outsrdeof the phragmocone;they haveno primary relationto the pro-ostracum. c. A problematicphragmoconeu,'ith a sheathand, continuing the latter in conical shape,a short rostrum; from the Lias e flower Toarcianlnear Hondelage(Braunschweig).The specimenbelongsto a private collection in Braunschweig;it was kindly provi{ed by its owner. The initial chamberis tentativelymarked in its presumedusual position. which is not distinct. The whole rostrum indeed merely forms an apex of the bro*'nish, glossysheathwhich becomesthinneranteriorly(nat. size).
lines. They are most distinct on the outer layer of the conotheca(ostracum) but they can also appear as
was described and illustrated (Fig. a7) as "Loligo
impressions on the hypostracumor, often slightly
aalensis and L. bollensls" by ZieIen (1830). Thus
stronger,on the periostracum.
Agassiz (1835) combined "Onychoteuthisprisca" wilh Belemnitesovaliss6and (169) called the resulting
The growth lines are of great importance for our knowledgeof belemnoidshells,sincethe shapeof the
especiallyto Belopeltis aalensis from the Lias, which
composite animal "Belemnosepia". Buckland had
free margin of the shell, especiallyof the pro-ostracum!
agreed
can be deduced from them. Buckland (1829) was
communications,but continued(1836) to distinguish
apparently the first to assume a continuation of the
the reconstructedbearerof the belemnitepro-ostracum
dorsal shell wall oppositeto the rostrum. Agassrzwas
from "Loligo" or "Sepiotelillrls" shells57,i.e. from our prototeuthoid (P1. 28-30) a fact which Agassiz
convinced by seeing such a structurein specrmens
with
Agassiz,
through
personal
fiom the collection of a Miss Philpott (p. 177, cf. Buckland 1836,Pl. 44', reproducedby Phillips 1867,
bitterly complainedabout (Transl.of Buckland, 1838,
PI. 8, Fig. 18). In fact there is no well preservedpro-
had recognized,like Agassiz, that Loligo shells were
ostracum,at the most some insignificantremains of its5.When studying the growth lines of the conotheca,
homologouswith the pro-ostracaof belemnites,but then (1836) also followed him in the confusion,
Voltz (1830,Figs 72,13) showedthat a tongue-shaped
althoughhe had recognized(p. 325) the insufficient
extensionof the dorsal shell margin had to be assumed.
congruenceof the gron'th lines. He consideredthis as a
Along with this insight an opportunityfor a long series
change occurring during growth and made his
see the explanationof Pl. 44'). Voltz (1836, p. 323)
of confusionswas provided. For the presumedpro-
subsequentreconstructionsaccordingly (cf. p. 109).
ostracaof belemnoidsshow an undeniablesimilarity to
The connection between Belopeltis aalensis and the
the shells ofthe prototeuthoids(p. 108) describedas "Onychoteuthis prisca" by Mrinster (1828), and
belemnite phragmoconewas in fact a forced one, rn that Voltz had changedthe characteristicgrowth lines
t04 Fig. 65. Some rare or problcmaticbelemnoids.o-i I lznat. size,
t.f\
r \
,t)M
\,-,,
k2l1nat.size. a. Phrogmoteuthisbisinuata. Phragmoconeand pro-ostracum. one lateral plate missing. Aftcr the original figure of E. Suess ( I 865).
$
A \/_,,,"----:\/
f/--'-7
b. Anterior part of a completepro-ostracum.l&rz. c. Dorsal view of phragmocone,drawn from the original 1/1. specimen,about d. Phragmocone of a belemnoid,afier Huxlcy (1864),in which Phragmoteuthls-likelateralplates could bc assumedto complete the pro-ostracum("dorsolateraland ventrolateralasymptotes"). (But compareFig. 63 b). e. "Belemnoteuthi,s"from the Lias of Lyme Regis "in slightly idealized representation"(Korschelt and Heidcr, Spec.part lll. p. 1144: "The figure is drawn liom a hithcrto un-described,very instructivespecimenfrom the collection of Dr. O. Jacckcl"). Probablythis is " A canthoteuthi,sconoca udci' (cf. p. 179). .l Septumof Diploconus belemnitoidesZittel from thc Tithonian (Strambergformation),after Zittel 1868. g. Fragment of phragrnoconeand sheath.idem. (v. thc fbrm of the suture drawn from the original specimen in the Munich
t,
collections). /r. Rostrumand phragmoconeafter Zittel.
,^. :^-.
l. Part of phragmoconein dorsalview. CompareFig. 73 a. where 'unrolled' only the conothecapresentsthis appearance.Median
\ i.)
and lateral plates here are retl narrow. k. "Conoreuthi,sdupiantts" after d'Orbigny (1842, Pl. 12) from the Lower Cretaceous. l. "Belemnoteullrl.rspec." after Langerhahn1906 (a problematic specimen"cf . Belentnoteuthis).
of B. aalensi,sso as to make them join the lines ol the
erroneous interpretations based on Belopelti.s shells
belemnite conotheca. He thus betrayed his own
were still published for some time. Thus d'Orbigny
p r i n c i p l e o f r e c o n s t r u c t i o n( F i g . 7 2 ) , p r o v o k i n g
(1842, Pi. 3, Fig. 3, PI. 4, Fig. 1) illustrates a
Quenstedt who had shown in 1839 that "Loligo bollensis is no belemniteorgan". But Qr.renstedt still
" Belemnites
aalensis"
by
simply
adding a
phragmocone with its sheath and rostrum to the gladius
consideredthe generally rounded posterior end of the
of Belopeltis aalensis (Zieten). In addition to gladii,
prototeuthoidshell to be the natural one, not realizing
tnre pro-ostraca have been described, however: Pearce
the necessityof assumingthe presenceof a conus (cf.
(1842, p. 185) indicated one for his new genus
p. 108).Voltz (1840) thereforereiteratedhis position.
Belemnoteutftis; Mantell (1848, Fig. 87) described the
For he recognized that the shells now called
pro-ostracum of B. (Cvlindroteuthis) attenuatus (:8.
"Belopeltis" are always incomplete in their posterior
puzosi d'Orb.), he writes: "This fossil comprises the
part. If one tries to completethem on the basis of the
following parts: L The capsule or periostracum. This
growth lines, one is oftenforced to assumethe former existenceofa conical structure(170) at the posterior end - which has subsequentlvbeen confirmed in
external investment, which consists of a thin, shelly, or
severalcases(p. I l4).
surrounds the
(171) The new idea of Voltz, i.e. the principle of
corno-calcareous integument that closely embraces the guard, and, (l 72) gradually enlarging upwards, finally
constituting
peristome
the thin
of
horny
the
phragmocone,
laminated
sheath or
r e c o n s t r u c t i o n sb a s e d o n g r o w t h l i n e s , w a s
receptacle, (that) has been described by all previous
subsequentlyaccepted by most authors, although
observers as an extension of what they termed the
105 sheathor capsule;within this receptaclethe ink-bag
accordingto Woodward it is "a horny dorsalpen, with
and otherviscerawere probablycontained...".A more detailed knowledge of the pro-ostracumcould not be
obscurelateral bands"). Finally Huxley observedin a species of Belemnoteuthisa "saddle-shaped"anterior
derived from these specimens,and the error made by Voltz (p. 168) could not be quickly rectified.euenstedt
margin of the pro-ostracum.(Probablya fragment).
of our "Acanthoteuthis"conocanda andspeciosa(p.
The existenceof a pro-ostracum as a continuation of the conotheca has since been confirmed by nlrmerous.finds; such pro-ostracahave been observed
180) must have had tongue-shapedpro-ostraca,but at the sametime gave his reasonsfor doubting that they
as isolatedparts (Fig. 63b) and in connectionwith the phragmocone (1). The former can be very well
were parts of a belernnite.He refusesto recognizethe apparently compressed phragmocones of
preseled if they have been rapidly buried after being
(1849) realizedthat belemnoidshells,especiallythose
Acanthotetrthisconocaudaas "alveoli", i.e. true (173) phragurocones (p. 530). If Miinster's specimenswere really identical with "Belemnites semisttlcatus" they would merely prove (p. 533) "that the shell in the belemnite alveolus did not have a circular margin but endedin a unilateralparabolicextensionthat cannotbe safely compared to loliginid shells". This at least admits, though reluctantly, the presenceof a proostracum;the belemnitesindeed were supposedto be
detached from the gas chambers (171) (Fig. 90), whereas the latter in most casesare badly damaged. With the only exceptions of Xiphoteuthis and Phragntoteulftis (Figs 66 and 67), the shapeis always that of a tongueas in Figures11,72 and73. As to the relative length, the growth lines do not necessarily provide a correct indication, becausethey cannot be easily followed in the hyperbolar zone, i.e. the lateral plates,where they are very closetogether.Great care is
closely related to the tetrabranchiatesand were not viewedas'hakedcephalopods".
therefore necessary,and isolated pro-ostracamust be taken into account as a complementarysource of information, especiallythe well preservedshells of
W o o d w a r d ( 18 5 I , M a n u a l ) a s s u m e st h a t t h e Belemnoideahad a pro-ostracum(of the type shown in
Acanthoteuthisspeciosass.The result is shown in Figure 90.
Fig.87). He correctly identifiesthe shells of '"Belemnosepia" as Geoteuthis Minst. (seeBelopeltis)
Specialattentionshould be given to Ihe connection
and placesthem in the "Teuthidae",which he contrasts with the Belemnitidae. Thus the general confusion may have been clarified. But Huxley (1864) describesa particularly
between the pro-ostracum and the conotheca; rr can seldom be observedin its three-dimensionalstate,but when it is, the similarity with the courseof the growth lines in isolatedphragmoconesis evident(Fig. 73). The insertion on the cone occupiesalmost the whole width
completebelernnitewith rostrum and ink sac (cf. Fig. 66c) from the Lias of Charmouth and curiously
of the latter. Here one can often obsere a third line in front of the last suture(Fig. 63f, double line 4 and 5),
considersit identical wtth BelentnosepiaAgassiz, i.e. Belopeltis (q. v.), which he views as a belemnitewith
which is particularly broad in adult belemnoids; this third line lies anteriorly to the last septum and -
pro-ostracumbut broken off phragmocone.fNote: Huxley's figure, copied by Naef in his Fig. 66c, was in fact of a compositespecimen].On the other hand, we
somewhat like a cross section marks the limit betweenconothecaand pro-ostracum(Fig. 906).It lies in fact behind the ventral shell margin and is due to the
owe him new, though partly vague, information on the
annulus (p. A); the latter, which had previously been observed only in nautiloids, thus also exists in belemnoids.- In this context we have to deal with the
formation of pro-ostraca,of which he distinguishes several types. First that of our Figure 87. It is "very thin and apparentlyhorny, or imperfectly calcified, in the dorsal region, and was supportedlaterally by two thin calcareousbands, or pillars, which inferiorly, expand upon the conotheca",adding here fHuxley,s] new genusXiphoteuthis (Fig. 66). Moreover, Huxley assumes that a third type should be found in Belemnoteutlrls Pearce(p. 185).(Accordingto pearce, the pro-ostracumof Belemnoteutliisis a cuttlebone,
questionof the morphologicalcharacter(homology) of the pro-ostracum;in other words we have to find out how the pro-ostracum relates to the general type of cephalopodshell (p. l4). An answerto this questionis provided by a straightforward comparison with an Orthoceras (Fig. l0). It thus appearsthat the ventral paft of the wall of the living chamberhas disappeared, as if it had been cut away, whereasthe dorsal wall has
106 relnainedin position.In termsof naturalprocesses this
also speakof an alveolarand a post-alveolarpart of the
n.reansthat the ventral wall atrophieddue to inhibition
"rostrum (.sensulato)". One rnay then distinguish
of its growth. The gap thus producedis closed,as in all
betweenmore cylindrical rostra and large and more
(p.22,Ft1. 10b),by Ihe muscularntantle. dibranchiates
club-shapedrostra with a small alveolar part, with
However, an alternativeexplanationis conceivable.
various transitionalforms (cf. Figs 71 and 95). The
(175) ln octopodsse there is no trace of a pro-ostracum
sheathaiso extendsin an anterior direction. as a thin
even in embryos. The shell rudiment is limited to the
envelope on the pro-ostracum, which may show
prospectiveposterior end of the mantle sac (Naef,
secondarysculpture unrelated to the primary growth
i 9 2 1 , C e p h a l o p o dvao, l . I I , P l . 2 5 , 3 3 a n d3 7 ) .I t s e e r n s
lines(cf. p. 105).
conceivablethat this could representthe primitive
The formation of a heavy periostracumis the result
condition in dibranchiates.The muscularmantle would
of its overgrowing the shell. While in orthocerathe
then have been closed dorsally like a barrel; at the
delicatejuvenile parts of the shell were simply cast off,
posteriorend one would have to assumethe presence
so that they would not be a continual source of
of an Orthoceras-like phragmoconecovered by the
disturbance due to inevitable breakage,
mantle skin, as has beenclaimed for Belemnoteuthis(q.
elimination is neither necessarynor possibleas soon as
v.). The pro-ostracumwould then have appearedas a
the shell fold is able to secretematerial covering the
secondary"protuberance"of the conothecareplacing
outside of the shell. Moreover, the periostracum
luch
the dorsal part of the muscularmantle. But this would
probably formed a v,eight addedto the posteriorend of
be a more complicatedexplanationthat is not justified
the body from early stagesonward (cf. Naef 1921,
as a simplerone suffices.
Cephalopoda, vol. I, p. I 10).
The shell partr so far discussedmust be considered as primarv, since their homologuesare seen in the
b) Materials for a general reconstruction of the
earliest nautiloids. In addition to these primary parts,
soft body.
a l l b e l e r n n o i d sh a v e s e c o n d a r y e l e m e n t s w h i c h
Belemnoidshellswith distinctremainsor impressions
collectivelyare called theperiostracum (p. 13). They
ofthe soft body are ratherrare (Figs 63 and 66). This is
lie on the outside of the ostracum and may show
easily explainedby its structureand by the occumence
different features in relation to the underlying
of the animals.Dead belemnoidsfloated on the sea
ostracum,or they may forrr a rather uniform envelope
surfacemuch like dead cuttlefish today, so they could
(Fig. 62). In any event. the periostracumis always
not possibly reach the sedimentintact to be buried in a
la,veredin such a way that conespackedone on top of
fresh state. Only a.fortunate coinc:idercecould cause
the other are formed: the innermostones are limited to
an intact animal to be cast ashoreand be buried there,
the region of the protoconch,whereasthe outermost
or to sink following rupture of the gas chambers,or to
extend to the free shell margin. The delicate.juvenile ,shellparts thus are most effectively protectedby the
suffocatein a shallow muddy basin (Solnhofen). The earliest,though uncertain,observationis again due to
periostracum,the subsequent,strongerones less and
Buckland (1829) who reportedon ink sacs from the
less so. In this way a "rostrum" appearsbehind the
EnglishLias, which he regardedas belongingto a co-
protoconch,i.e. a more or less pointed or massive
occumingbelemnite(8. ovalis)(cf. Br.rckland 1836,Pl.
development of the periostracum surrounding the
44' , Ftg. 7). His observationswere confinned by
conotheca. We call the alveolar part of the
A g a s s i z ( 1 8 3 5 ) u , h o ( a c c o r d i n gt o B u c k l a n d 1 8 3 6 ,
periostracum the "sheath" sensLtstt,icto,the post-
Jahrb.P. 38) "(working throughthe colle-ctions of Miss
alveolar part the "rostrum" s. str. In the literature we
Philpots at Lyme-Regisin Octobcr l.q3-l)discovered ( 1 7 ' 1 t w o i m p o r t a n tv. e r y i n s l r t rrei r e s p e c i m e n si n.
find very hazy concepts: Since the thickenedshell wall, togetherwith the
which the ink sac was still in placc in the anterior
rostrum proper, forms a solid mass which nearly
h o r n y s h e a t h o f a n i n t a c t b c l r - n t n r t e .a n d w h o
aiways forms the greater part or whole of a well
henceforthintendsto placeall bc-lc'nlritc-s in one genus
preservedfossil, and since in most reconstructions of
of the classCephalopoda".tbr shrch hc proposedthe
completebelemnoids(I 76) Ihis solid structureappears
nameBelemnosepia(p. 169\.
as a homogeneoLts, very distinctoccessory,one may
O w e n ( 1 8 4 7 ,P h i l . T r a n s. l . l j r . i l s of o u n d a n i n k
r07 sacln Belemnotelrthisantiqua (q. v.) in the Middle Jurassicof Christian Malford (:"Belemnites owenii"
Mtinster was apparentlythe first to observe these fossils;he called them "OnychoteuthisLichtenstein",
, . 5 3 5 ,P l . 3 6 , F i g s4 , 5 , J , 9 , l 3 ) . I n Q u e n s t e d1t8 4 9 p several casesthe phragmoconeswere broken so that
since he consideredthem to be members of a recent type of oegopsidbearing hooks. But he also confused
the ink sacs were lying in the last chambers (cf.
them with several other types in which he merely
Quenstedt,p. 530). Overall thesefossils were not very well preserved. H. v. Meyer (1832,p. 322)had already
assumedthe presenceofhooks; thereforewe cannotbe
reported on a beiemnite with an ink sac "at the upper
certain today what he really meant to include when refeningto Onychoteuthis prisca (1828,p. 581) and O.
end". This specimen came from the Lias of Banz (Swabia[Bavaria]), and the authorassumedthe general
angusta (1830,p. 404, 458). At any rate, several prototeuthoidswhich are in fact devoid of hooks were
occurrenceof an ink sac in belemnites.The same rs
included6o (cf. p. 122).
believed by Buckland (1836, Jahrb.,p. 39-40) who takes it as evidencefor an internal shell, "for the ink sac replaces the protective shell in the naked
Diagnosis:Acanthoteuthlsdesignatesbelemnoids of which the rostrum is not known with cerlainty,while the phragmoconeand pro-ostracum are like those of
c e p h a l o p o d st c" t .p . 2 4 1 .
Belemnitidae and the arms each bear two rows of
Suryrisingly no well preservedbeaksof belemnites have been found, and distinct remainswere only found
hooksas in Belemnitidae.
l a t e r( c f . F i g . 6 6 ) , s o V o l t z ( 1 8 3 0 ,p . 3 3 ) a s s e r t e tdh e absence of such parts in both ammonites and
The following speciesmust be consideredstill valid:
belemnites. The "genuslcanthoteuthis" Wagner 1832[1S39]. The evidencefor an ink sac in various belemnoids demonstratedtheir dibranchiatenature.Early on much more extensive knowledge of these animals was acquired, but unfortunately it was impossible to determine the affinities of the fossils in question.For the identificationof belemnoid speciesis in general
l. AcsnthoteuthismonteJioreiBuckman I 880. Here bclongsBelemnoteuthis montet'iorei Buckman1880 (Proc.Dorset.Nat. Hist.,and Antiqu.Field Club,vo1.3, p. 141)andCrick(1902,Pl. 1). This is an incompletely preservedbelemnoidbody from the Lower Lias between Lyme Regis and Charmouth,showing a gross outline of the mantle sac, a very large ink sac and five to six distinct arms with
basedon the rostrum; but in most casesthe rostrum is
double rows of hooks. fNote: Buckman's figured specimenis in fact composite].(179) Two of these
missing in the best preserved animal bodies and phragmoconeswith a pro-ostracum(p. 170). If it is
than the others. All of them bear very peculiar hooks
present, we are able to assign the specimensto the f a m i l i e s B e le m n o t e u t h i d a e ( q . v . ! c f . a l s o
that fiudging from the original figure) have the shape shown in our Figure 689; they are orientatedat right
Phragntoteuthis)or Belemnitidae(Fig. 67) (perhaps
angles to the axis of the arm. Apparently they were involuntarily retracted(like the claws of a cat) before
further types will be found). If the rostrum is missing, identification remains (178) uncertain,since other features are lacking, i.e. are not recognized as such (hooks,Fig. 68!?). For indeterminatebelemnoid bodies and shells without a rostrum, we therefore use the lictitious generic name proposedby R. Wagner; it is all the more necessaryas these fossils are of great interest.This interest derives from the presenceof hook-bearing arms, in which we have recognized(p. 26-30)proof of the decapodcharacterofthe belemnoids(cf. Fig. 91, a n dp . 1 8 1 ) .
arms are markedly shofter, two appearmarkedly longer
striking, and after death this position was preserved. Crick considersthis animal to be a belemnite without giving his reasons.(Also cf. Crick 1907). 2. AcanthoteuthisconocazdaQuenstedt1849. Herebelong:Onychoteuthis prr.rcaMiinst. 1828,p. 581 (in part?).Onychoteuthis prisca Meyer 1832,p. 322 (in paft?). A c o n t h o t e u t h i sp r i s c a V o l t z 1 8 3 5 ,p . I ( i n p a r t ? ) . Onyc'hoteuthis conoc.cttrda 1849,p. 529,550.556; Quenstedt Pl. 36,Figs6-8,12,14.Onychoteuthis conocauda ibid.1858. p.245. T h i s s p e c i e s i s r e p r e s e n t e db y c o m p r e s s e d
108 phragmocones,often associatedwith remains of the
(crochets)ofthis cephalopodscatteredall around.Both
pro-ostracum,ink sac,muscularmantle,head and arms
objectslie so closeto one another,partly overlapping.
(with double rows of hooks),as well as individual parts
that one might believe that they belong to one and the
from the above list, from the Lias e fl-ower Toarcian] (black "Tafelfleinz") of Swabia (and England); see the
sarneanimal, but closer examinationrevealsthat they representtwo different animals, namely Belemnites
original figures by Quenstedt.Apparently these are belemnites with a very short rostrum like those
semistrlcatus atd Onychoteuthisspeciosa(the largest
occurring in the same strata (-8. incurvatus Ziet.?)
Buckland'sBelemno,sepia in the Liassic shalesand in
which
similarly cornpressed
the lithographic stone were in vain; in no German
p h r a g m o c o n e .( l o c a l i t i e s : H o l z m a d e n , P l i e n s b a c h ,
collectionknown to me did I find a true Belemnosepia,
Banz).
often show a
M i i n s t e r( 1 8 2 8 ,p . 5 8 1 ) p r o b a b l yb a s e dh i s
"Onychoteuthisprisca" on specimensof this species rather than on Belopeltis aalensis (Fig. 47); otherwise the namewould be unintellieible.
fossil speciesknown to me). All my efforts to find
which I first suspectedto be representedby the above mentionedbody. Miinster's doubts were apparentlymisplaced.Here we have a single animal lacking the rostrum; only the latter could prove affiliationwith B. semisulcatrs.The
3. Acanthoteuthisjaeckeli n. sp. (?).
hollow continuationof the phragmocone unchambered,
Herebelongs:BelemnoteLrthi,s spec.JaeckclfJaekel]1890.p. 92.Belemnotezrllzls spcc.KorschcltandHeider1893,vol. III, p. 1144.Fig.679.(Perhaps theprevious species!).
of courseis the pro-ostracum,(181) whrch was not yet
The original figure of this very fine specimen(Fig.
speaks of an "exceptionally large specin.renof B.
fully understoodby Miinster. W a g n e r( 1 8 6 0 ,p . 2 9 ) m e n t i o n st h e s a m ef o s s i l .H e
65e) from Lyme Regis is probably a reconstruction.
with a large phragtrocone,body, head semisulcatu,s"
Whetherthe outline of the pro-ostracumis really intact
and individual hooks next to the latter.The shapeofthe
remainsvery doubtful.At any rate,it doesnot resemble that of normal belemnoids(Fig. 73) which can also be
body is said to be as in Ac. .ferussacli.Ren-rainsof the "brown, horny, irregularly furrowed" pro-ostracumare
assumedfor these forms (to judge frorn indistinct
mentioned.Accordingto pages72-73 of this work the
growth lines seen on the very thin, glossy pro-ostraca
rostrum is lacking. The description is misleading,
of Ac. c'onocauda).
however,and thereforeneedsto be mentioned. Here also: Acanthoteuthisspeciosa,./brrussat'ii,
(180)
lichtensteiniiMrinst. 1839(p. 105,Pl. 9 and 10,Fig. 14. AcanthoteuthisspeciosaMiinst. 18396r.
2). These three fossils probably representthe same
Part of "Belemnitessemisulcatus"Miinst. 1830 (p. 7,
s p e c i e s ,a s a l r e a d ys u p p o s e db y d ' O r b i g n y ( 1 8 3 0 ) ;
P l . l , F i g s 1 , 8 . 1 5 ) a l s o b e l o n g sh e r e . ( S e e a l s o
Miinster maintainedhis distinctionof three species.All
B u c k l a n d 1 8 3 6 , P l . 4 4 ' ) . P h r a g m o c o n e sa n d p r o -
are basedon bodieswith arrns.It is true that d'Orbigny
ostracaof this form belong here as long as association
erroneouslyrelatedthem to Plesioteuthispriscu (1.842,
with a rostrum is unknown or uncertain.In contrast,the
P a l . f r . j u r . , P l . 2 3 , F i g . 2 - 4 a n d 1 8 4 5 ,p . 4 0 7 , P l . 2 8 ) ,
identity of this kind of shell (devoid of a rostrum)with
partly under the name Celaenoprisca, partly under the
bodieslike those shown in Figs 639 and 91, i.e. with Ac. speciosaMtinst., as proposedby Zittel, must be
present name. (The first one is from Miinster's manuscriptof 1836)(cf. p. 1l5). - Quenstedt( 1849,Pl.
consideredcorrect,especiallyon accountof a specimen
36, Fig. I 1, p. 532-533)rightly recognizedthe typical
in Munich (exhibited collection), namely Zittel's
similarity wtth Ac. ("On1:choteuthis") conocaudabut
O s t r a c o t e u t h i s s t r p e r b a ( p . 5 1 0 - 5I 1 ) , w h i c h w a s
assumeda clear differencefrom belemnites(p. 173).
known to Miinster. The latter apparently understood
Morris (1854) placedAc. speciosain Belemnoteuthis.
the connection. He writes(1836,Jahrb.,letterto Bronn,
Wagner (1860) restricted the name to those hook-
p. 583):
b e a r i n g c e p h a l o p o d so f t h e J u r a s s i c w h i c h h e
"From SolnhofenI got the large alveoiar cone of a belemnite with the unchambered,hollow continuation
consideredas "loligineans"or "teuthodans". Following theseobservations, basedon Miinster's
of the shell, besidewhich lies the damagedsac of a
s p e c i m e n si n t h e M u n i c h c o l l e c t i o n s ,a c o m p l e t e
very large Onychoteutlrls, with small arm hooks
descriptionof the phragmoconeand pro-ostracum(Fig.
t09 90), and an overall picture of the mantle sac, head and
contrast,clear differentiationof the tentacular(183)
arms can be achievedon the basis of the rich material
arms did not exist. In this respectAc. speciosaappears
from the Upper Jurassic of Solnhofen, Eichstiitt,
closeto Belemnoteutirls;however, it may rnerely show
Daiting and Nusplingen. The phragmoconeshows all
the typical condition in all later belemnoids,which is
the featurestypical of a belemnite(cf. Zittel 1885,p.
different from that in other decapodgroups. We may
511). The same can be said for the pro-ostracum,
ask ourselveswhether an inconspicuousdifferentiation
which is perfectly well preservedin a specimenfrom Solnhofen, now housed in Munich (Fig. 90).
of the fourth arm pair (counted frotr above) is neverthelesslikely. As typical tentaculararms exist in
Comparablespecimensshow slight variations: a very
both sepioids and teuthoids, two groups whose
narrow, flat median keel may be recognizableand the
separationcan be datedback at leastto the Lower Lias.
accompanyingribs may become broader or narrower.
one can indeedassumethat the condition for a division
The marginal zone apparently was delicate and
of labour between arm pairs was already attained in Triassicdecapods.It seemsconceivablethat a certain
uncalcified. (182) The mantle.sacapparentlyhas the typical form, but is never well preserved. Traces of the
adjustmentin responseto ecologicalconditionstook place in later belemnoids,e.g. with availability of prey
muscular mantle show transversestriation like that seen in teuthoids (Fig. a8). An ink sac is often
animalsthat were powerful rather than fast swimming,
recognizable(Fig. 63e, g). I never lound clear tracesof
(oceanic forms), in which either the tentaclesare lost
the fins. In teuthoids,which were more rapidly buried (p. 176) conditions for their preservationwere of
during late ontogeny (p. 27), or they become increasinglysimilar to the other arms during post-
course more favourable (Fig. 42). Likewise the head,
embryonic development (Ommatostrephessagittatus
eyes and beakshave only left indistinct impressions,
(Lam.)(cf. Fig. 62).
similar to what we observe in certain metateuthoids
which neverthelessconfirm the overall picture (Fig. 62). The reconstruction of the whole animal correspondsto Figure 67e. Here Mrinster's, Wagner's and Zittel's Solnhofenspecimen,which was mentioned above(p. 180), is of great importanceas it showsthe phragmocone,pro-ostracum.mantle sac,headand arms together, thus providing good information about the relative sizesof theseparts.In parlicular,the respective proportionsof the soft parts and the shell can only be assessedfrom this single frnd, whereasmore complete fossils are available for a study of details of the different parts. (For an assessmentof the overall aspect,see also the
somewhatindistinct - picture of
5. Acanthoteuthis problematica n. sp. This is a fossilfrom Daiting(lithographic limestones. Upper Malm).Slabandcounteryart arehousedin Munichandhave neverbeenproperlydescribed, althoughthey reveala very peculiaranimalwhich lookslikc no other(Fig. 56e).I have beenconfusedby the previousliteraturewhich spoiledmy characterization ofa majorgroup(1921,Cephalopoda, vol. I, p. 147underCelaeno). Wagner(1860,p. 35) identifiedand describedthis peculiar form as a representativeof his new Celaeno conica.ln doing so he confirmedbelemnoidfeaturesin a teuthoid, especiallythe occurrenceof hooks, which
the fine specimenin the frontispieceof Abel's 1916
was erroneously(at any rate without justification)
book. For a specimen with an ostensible "buccal
assumed for fossil teuthoids. This idea haunted Miinster (1828), Meyer (1832), d'Orbigny (1842,
membrane",seeCrick 1900). The arms, shown in Figure 91, are of special
1845) and subsequentauthorsand thus becamepart of
interest. There must have been 10 arms of somewhat
the literature. Since Wagner's indications were more
variablelength and thickness62, but of sirnilar structure.
recentlyconfirmedby J. Walter (1905),I took it as a confirmed fact and assumedthat the transformationof
Each arm bears two rows of hooks6rthat are very similar in shapeto those of Belemnoteuthis(Fig.68e), and clearly different from those of the Acanthoteuthis speciesof the Lias. Whetherthere were suckersas well as hooks (p. 29) cannotbe ascerlained. The dorsal and ventral arms were probably weaker than the lateralarms, as in many recentdecapods;in
part of the suckersinto hooks was a primary teuthoid feature derived (184) from the belemnoids (Cephalopoda,vol. I, p. 127-132);this was a major error. Wagner's Celaeno conica (p. 151) is a very strange fossil, perhapsrepresentinga new genus and family, that conveys the impressionof a belemnoid
110 Fig. 66. The reconstructionofbelemnoids. a. Xiphoteuthiselongata,shell in lateralview. &. The same. in ventral view with mantle sac added. Reconstructedfrom the original figures ofHuxley (1864). 1. dorsal projection, 2. ventral projection of mantle margin. 3. pro-ostracum. 4. thin part of thc lattcr. J. mantlc sac. 6. insertion of muscular mantle on the shell margin (7), 8. 2/5nat. conotheca.9. sheath,10. rostrum. size. c. Belemnites brtrghieri (p. 111) from the Upper Lias of England. Shell with soft parts on a shale s1ab./r. hooks, a. arms, fr. mandible, v. anterior margin of pro-ostracurn.p/i. phragmocone.r. rostrum. l/2 nat. size. As far as I know. this is the only specimenwhich showsthe shell with the rostrum and pro-ostracumtogctherwith thc head and arms.Although thc statcofpreservationis not very good as to the details,thc
\
picture neverthelessconfirms our generalassumptionsrather
I
nicely (Figs 63 and 67).
I
I \
t
imitation of the teuthoid Celaeno,so to speak.lt shows
\
(Fig. 56a, d) the head, arms with hooks, the mantle
I
sac,and the conical shell. The latter is evidentlyplaced with the opening down and then compressed,which resulted in the form of Celaeno. Wagner did not
/
II
)1
realize that such deformation could have occurred either during burial or due to a thorough rearrangement of the belemnoid type; he merely saw a cephalopod with the posteriorpart in the fonn of a flat cone which is so characteristicof the Celaenidae.For the rest. he givesa gooddescription: "The head is of moderate size, the mantle sac rs extendedto where the shell disk, which is no longer present6l,begins; it ends with a broadly rounded posterior part. Even though the substanceof the disk has disappeared,it is still traceableas an impression, oval in outline, deeply concave in the middle, and traversedby several delicate, concentric, oval ringst'5 that are parallel to the margin of the disk, in other words showing the essentialfeaturesof the disk of C. conica; I therefore interpret this specimen as an imprint of this species. Particularlyremarkableare the arms, which are closely packed, having lost most of their distal ends.They are equippedwith numerous small hooks66similar to Acanthoteuthisfbnrssacii; however, in several arms one also finds longitudinai seriesof rings with hollow centres(185), which given their shapeand position- must be impressions of suckers".
t 1 1
t l r
(( b )r:
V"
Fig. 67. - Reconstructions of somebelemnoids. o. Belemnotetrthis antiqua'12nat. size,after the original figuresof Suess( 1865). D.Crosssectionofposterior parl ofcone. c. Phragmoteuthisbisinuata.r/2nat. size. d. BelentnitesgiganteusSchloth.I 'nnat. size (uvenile form; p. 239) e. The same(cf. figurep. 211).
The genusBelemnoteuthis.
fins are added,following our generalprinciple (p. 7). Considering the observed structures and their
The ideasbasedon the animalsof "Acanthoteuthis"are
necessaryaccompaniments, we find almost complete
further reinforced by observations on a particularly
agreementwith the data on "Acanthoteuthis" speciosa
well preservedgenus of belemnoid decapodfrom the
(p. 180 and 111). In spite of this and of the
Upper Dogger flate Middle Jurassic] of Christian
stratigraphiccoincidence,and the similar shapeof the
Malford, which has been publishedby Pearce(1842)
hooks (p. 189),we cannotunite theseforms, because
and Owen (1844). The specimensshow the typical
Belemnoteutirlsshowsa well preservedsheathof quite
arrangementand proportions (Fig. 67a, b) of the
specific character(cf. relevant chapter).On the other
phragmocone,mantle sac,head with eyes and arms, so
hand,we do not find clear evidencefor the allegedlack
that the overall aspect of the best exarnples(186.)
of a pro-ostracum,althoughnothing is known about its
providesa good illustration of the anirnal.In the figure
propefties(cf. Fig. 90).
only the outlines are complete in their details and the
( 1 8 7 ) O u r i d e a s o f t h e t y p i c a l c h a r a c t e r so f
r12 Fig, 68. -Morphology ofhooks in fossil and recentdecapods. problentaticrzfrom Daiting (p. 151).20/1. a. Ac'anthoteuthis r/1. b. Acanthoteuthisspeciosofrom Solnhofen(p. 180). 1/2. c. "Onychites"spec.after Quenstedt1885. 3/y. cl.Phragntoteullrlsfrom the Triassicafter Mojsisowicz. 7/1. e. Belenlnoteuthis cmtiquaalter Pearce1854.
f,C f
v;.
r@.
['
f,
.f. "Onvchites"spec.from the Middle Jurassic,after Quenstedt 1g95.2/r. 7/1. g. Belemnoteuthis ntontefiorelaftcr Crick. h. Acanthoteuthisc'onoc'auda from the Lias e flower Toarcian]of Holzmaden.6/1. i. Onychitesspec.from Nusplingen(Munich collections).2/1. /r-o.Tentacularclub of a youngAncistt'oteuthislichtenslelnl(Naet. Cephalopoda, vol. 1. p. 13 1). i. sucker(r in o) with strong.hook-like tooth (112). /. young hook. somewhatmore developedbut still showing the
[,
(
sucker. n. small,fully developedhook. r. large,fully developedhook, viewed from belor,v. l o l ec l u b .r 1 n a l .s i z e . o . r .h p. large hook in latcral vicw. Kp. hood;ilft. hook; Sl. stem of hook; Hr. homy ring of sucker;R:. adhesivering of suckerhorny ring; Rd. marginalring; Ba. basal part of modified horny ring; El. grooveson it (impressionsof blood vessels);Fo. extensionon it; Ir. suckercamieror hook carrier;1. distalhook-likesuckersimilarto /r (cf. Kef-erstein 1866,Pl. l3l).
belemnoid decapodsappearrather vaguely confirmed
grainedshales.
by fairly well preserved belemnite animals (with
We have alreadydiscussedthe generalmorphology
rostrum) occurringin the English Lias (cf. Fig. 66c).
of hooks (p. 29) and now wish to apply that
( S e ea l s o K e f e r s t e i n 1, 8 6 6 ,P l . 1 3 1 ,F i g . 8 , a n d Z i t t e l
knowledge. Let us first look at the special aspectsof
1 8 8 5 ,p . 4 9 8 , F i g . 6 8 1 , G r u n d z i i g e1 9 2 1 ,p . 5 8 5 , F i g .
belemnoid hooks: these structuresare not apparently
1256; these figures are potentially confusing due to
identical with those of recent teuthoids (which again
arbitrarilydrawn outlines).
show great differences among themselves) and therefore cannot be easily interpreted. Interpretation
As far as arms are sufficiently well preservedin the
was not even attemptedby Quenstedt(1858, p. 201) who was the first to pay special attention to these
f o s s i l r e c o r d o f b e l e m n o i d s ,b e g i n n i n g w i t h t h e
structures,calling them "onychites".He distinguished
T r i a s s i c P h r a g m o t e u t h i s , a n d c o n t i n u i n gw i t h
(1885,p. 512) differentspecies(e.9.O. ornatLrs, uncus,
Belemnotettthis, Acanthoteuthis and Belemnites,these
c) The brachial armament of Belemnoidea.
arms bear structures resembling the hooks of recent
runcinattts) fron' the Lias, Dogger and Malm. The oldest ones known to hirn were from Lias y (O.
teuthoids;they can only be interpretedin analogyto the
numismalis).As regardssome isolatedspecimensthat
latter(Fig. 68). I havenowherefound (p. 184)reliable
he placedhere, doubtsremain as to whetherthey really
tracesof suckers(not even in the teuthoids,which must
were from cephalopods(Fig. 68f, i); one cannot easily
have posessedsome). Apparently these structures,
understandhow they could fit in and what their
despitetheir "horny rings" (p. 27), were too delicateto
function may have been. In type i the terminal point
be preservedin fossils.In the hooks,however,the stem
was apparentlylost, so that the whole element is not
at least was strong enoughto becomefossilized;the
c l e a r l y h o o k - s h a p e dT. h e f o r m s a , b , d , € , g , h ,
points can only be observed in exceptionally fine
however,must have been belemnoidhooks, since they
r 13 have been found in association(188) wtth identifiable
PassaloteLtthls)shows a clearly different hook shape
animal bodies; they differ from the hooks of recent
(Huxley 1864) from Ac. specio,sa. But a comparative
teuthoidsin that the proximal end is ratherpointed and
analysisof the utility of these elementscan only be
only distally grades into a thicker part that may be
expectedfrom a new study of richer material.
interpretedas the remains of a horny ring. There rs no broadened"root" to the hook. Perhapsit was softer
d) On the position of the shell inside the soft body
than the shaft of the hook proper and thus was
of the belemnoidanimal.
destroyedduring fossilization(?).The curved end is
The fossils discussedso far may provide us with some
often missing altogether,so that the hooks may have
generalideas,but they give us no detailedinforrnation
lookedmore iike spines.
as to the relationshipof belemnoid shells to their soft
The occurrenceof these structures.which in the
parts. To acquire such detailed knowledge we need
Middie Jurassic attain the size of a little finger (Megateuthis?)is widely observed.The basic forms are
comparison with recent forms. Voltz already recognizedthis requirement.In addition to clarifying
observedtogetherwith cerlain shellsand soft parts (see
the principle of shell growth (p. 168)6?, he was also a
above!).Moreover,they often occur in isolation(Lias,
pioneer in investigatingthe close relationshipbetween
D o g g e r , M a l m ) ( T r i a s s i c ? ) . S o m e t i m e st h e y a r e
belemnitesand recent cephalopods,(190l looking first
obserr,red in coprolites,i.e. in concentrationsof organic
for general correspondences(1830), and later for
remains,especiallyin the Upper Jurassiclithographic
specialconformities.He found the latter(1835,p. 5.1in
limestones.Miinster found them togetherwith remains
the ommatostrephids(."Loligo sagittata") (cf. Fig. 59),
of the gladius (pro-ostracum) in the stomachs of
but he also attemptedto derive all dibranchiateshells
Plesioteuthis, and curiously enough he took this as
from the belemniteshell, especia[lyhis "Onychoteuthis
evidence of the occurrenceof hooks in Plesioteuthis
prisca"
itself. In the stomachsof ichthyosaurians, massesof
Prototeuthoidea,p. 178), "Loligo sagittato" (i.e.
hooks (and belemnite rostra) are found, thus
p. 158),Loligo vttlgaris(i.e.Loliginidae,p. Oegopsida,
confirming their generaloccurrencein belemnites.We
1 5 8 ) , " T e u d o p s i s " ( i . e . M e s o t e u t h o i d e ap, . 1 3 5 ) ,
have no clear evidence for the earliest forms of
"Onltchotetrthisangusta"(i.e.Plesioteuthis prisca, p.
belemnoidsin the Triassic: it is unknown (Fig. 62) whether they bore hooks, and if so, whether this
ll4), "Octopas", i.e. Octopodidae,the shell rudiments o f w h i c h h e d i s c o v e r e (d1 8 3 5 ,p . 1 ) . - H i s m e t h o do f
occurredon all the arms, and whether such occurrence
homologising is still partly valid today. Only the
was uniform. As there is no good evidencefor the
relationship with the sepioid decapods was not
occurrenceof hooks in the Aulacoceratidae,and since
recognizedin detail by Voltz. I have now been able to
the older (fossil) representatives ofthe teuthoidswhich
show this relationship(Part II). D'Orbigny foilowed
haveto be derivedfrom the oldestbelemnoids(p. 161)
Voltz in his ideas;he was especiallyimpressedby the
lack any trace of hooks (incidentally, they are totally
similarity of belemniteand ommatostrephidgladii (e.g.
lacking in all the sepioids),it seemslikely that the
1 8 3 9 ,p . X X X V ) . ( S e ea l s o 1 8 4 2 :A n n . S c .N . , p . 3 6 6 ) .
transfbrmationof part of the suckersoccurredonly in
B u c k l a n d( 1 8 3 6 ,P l . 4 4 ' , F i g . 2 ) h a d w h o l l y e n o n e o u s
two groups(later belemnoids,oegopsids);althoughthis
ideas about the insertion of belemnoid shells into the
transformationis foreshadowedin the decapodsucker (p.27), it shouldnot be assumedto be characteristicfor
animal (cf. below, Fig. 76b). His reconstructionshows
the generaltype of the group, which of coursewas of
phragmoconeinto the mantlesac.
(cf. p. 165). belemnoidcharacter For speciesdistinctions,the "onychites" provide
(i.e. the
erroneously interpreted
a sepioid character with deep penetration of the We considerthe following factsessential: l.ln
all well preservedbelemnoids the
reliable indications,as much as for the relationshipof
phragmoconeoccupiesthe whole end of the
certain types. The agreementof hook shapesbetween
mantle sac and continues it, as it were. This
A c a n t h o t e u t h i s s p e c i o s a a n dA c a n t h o t e u t h i s
suggeststhat the insertion of the muscular
monte/iorei,and the differencebetweenAcanthoteuthis
mantle followed the free shell margin, not
ntontefiorei andA. conocauda.is clearly recognizable
only of the phragmoconebut also along the
in Figure 68. (189) B. brughiet'l in its turn 1cf.
pro-ostracum (Figs63, 66,61).
t14 2.In fossil prototeuthoidsstudiedwith the above
o f t h e p h r a g m o c o n e .N e v e r t h e l e s st h e c e n t r e o f
observationin mind, the assumption is
buoyancy is situated far back in the animal so that a
confirmed. This form of insertion can be seen
horizontal swimming position is difficult to stabilize.
in Plesioteutftis(Fig. 42) in particular.Here,
Without compensation(p. 192) such a position is
furthermore,it extendsto the inner side of the
possibleonly at the seasurface.
margin of the pro-ostracum. 3 .R e c e n t t e u t h o i d s a l w a y s
(192) 3. D'Orbigny supposedthe function of the show
the
rostrum to be protective.According to him it is a
o n t o g e n e t i c a l l yp r i m a r y i n s e r t i o n o f t h e
"protective device" against "shocks" and a defensive
muscular mantle on the free margin of the
organ6e.We have seen (p. 176) that it forms a
shell, even though secondaryshifts during
necessarycorrectiveitem in internal shells.D'Orbigny
post-embryonic development may be far-
already recognized that the buoyancy of the
reaching(Naef 1923,(191) Cephalopoda, vol.
phragmoconemust be compensatedfor by the rostrum.
I, chapter5). If the conus is well developed.
This of courseis the more effectiveas the weight of the
the primary inserlionis conservedin this area.
rostrum increasesand as its weight is placed more
4. ln older embryos and very young "larvae" of
posteriorly(club shape in Atractites, Hastitinae,
oegopsidteuthoids,the whole shell is inserted
Belemnopsina).Increasingthe weight of the alveolar
in a "belemnoid" fashion (Fig. 61a), in that
part in the areaof the large gas chamberswould merely
the insertion of the muscular mantle strictly
compensatefor their buoyancy. If however the centre
follows the free margin of the shell.
of gravity lies far behind the effective center of it and thus allows the buoyancy, it counterbalances
Thus the way that belemnoid shells are insertedin the
animal to maintain a horizontal orientation when
muscular mantle, as suggestedby well preserved
swimrning in midwater. Therefore forms with a long
r e m a i n s( F i g .6 7 b ) .i s n i c e l yc o n f i r m e d .
rostrummust havebeenparticularlygood swimmers. Although there are many differencesin detail, the effects of which should not be underestimated,the
C. On the function of typical belemnoid shells
above considerationsneverthelessallow us to form
and the life stvle of their bearers.
some general ideas on the life style of belemnoids. Indeed, the main elementsof their organisationare
The function of the internal shells of dibranchiate
essentiallythe same:belemnoidsare slenderdecapods
cephalopods was already surveyed by d'Orbigny
with arm hooks and a conical, gas-filledposterior end,
( 1 8 4 2 , p . 3 6 8 ) w h o c a r e f u l l y c o n s i d e r e dt h e
the most conspicuousdifferencesbeing observedin the
morphological features of the different parts of the
shapeof the terminal projection, the mass of which is
shell.Clearly they do not haveidenticalfunctions.
relatively insignificant.We regardthem all as nektonic
1. The function of the pro-ostracum("lame
forms of surface waters and coastal zones, only
cornde") is most readily assessed,since it persists
specializedforms having diverged to live in deeper
largely unchangedin the teuthoidsas a "gladius" which
parts of the sea and in the open sea70.The mass
is easily observable.It acts like the backbone in
occurrenceof many speciessuggestsa gregariouslife
vertebrates"to support the flesh". (In forms where it has considerable width, it rnay also passivelyassistin
style similar to that of the [geologically] younger teuthoids; we see no reason to assume essential
locomotion in that its elasticity helps the mantle to
differencesfrom the life style of the latter, as far as the
expand after each muscular contraction.Where it is
heaviershellpermitted.
(Fig. 59), this narrow, as in the Ommatostrephidae
The belemnoidsin general7lcan by no means be
action is compensatedfor by muscular development).
considered as creeping, (193) benlhic forms, and
(SeeNaef, Cephalopoda, vol. I, chapters5 and 32).
observationssuggestingsuch a life style are erroneous.
2. The gas chamber.rhave the same effect as the
The "track" of Acanthoteuthis desuibed by Jaeckel
swim bladder in a fish, although funlike the swrm bladder]they cannotchangetheir volumes68. It must be
201) must be interpreted differently: in the platy
recalledthat the largestchamberslie at the anteriorend
limestonesof Solnhofenone finds the same imorint
( 1 8 9 9 ,Z . d . d . g e o l .G e s . ,p . 3 6 ) a n d W a l t e r( 1 9 0 4 ,p .
115 repeatedat a small distance.This is probably a seriesof
affiliation often being questionable.
imprints of an apparently stiff arm crown (probably
Belemnite rostra have been found since ancienl
with rigor mortis) of Acanthoteuthis which were produced one next to the other laterally, entirely
times. The name Belemnltesis due to Agricola (1546).
automatically,i.e. in a physiologicallyinconceivable
niger" shouid be placed in the genusPassaloteuthis;tt
condition.The original slabs can be analysedin the
has often, probably erroneously,been identifiedwith B.
Munich collections.Moreover one shouldrecall that no
paxillosus Schloth.
Lister (1678) first used it as a genericname.His "B.
k n o w n l i v i n g t e u t h o i d r 2s e t t l e s o n t h e b o t t o m ,
The interpretationsof thesecommon fossils which
especiallyon muddy bottoms; the hook-bearingfbrms
were given in ancient times are not of scientilic
in particularare offshoreand deepseaswimmers.
importance.They were said to be "thunder bolts",
Nutrition in belemnoids also must have been
a m b e r p l u g s , s t a l a c t i t e s , s e a - c u c u m b e r s t, h o r n y
similar to that in recent and fossil teuthoids.As in the
a p p e n d i c e s ,m a m m a l o r f i s h t e e t h , " s e a t u b e s " ,
latter, faecesdemonstratethe existenceof cannibalism.
solidified urine of the lynx (penis bone!) etc. Ehrhardt
They were doubtlesspurely carnivorouspredators.
(1724) was apparently the first to recognize the
Fish, crustaceansand their relatives are the matn prev
relationship of the phragmocone to nautilids and
ofall dibranchiate (seep. I 16). cephalopods
ammonites.The relationshrpto Sepia (the rostrum)was n o t a s e v i d e n t . I t w a s n e v e r t h e l e s sa s s e r t e db y T h e o p h r a s t u sa,n d B l a i n v i l l e ( 1 8 2 7 ) l i s t s 8 1 e a r l i e r
D. The family Belemnitidae (d'Orb. 1845)s. restr.
authors as having confirmed this opinion. There were not many good reasonsfor it, so this opinion cannotbe
Content,t'. L Gencralaspects. A. Prcliminary remarks(below). B . D i a g n o s i s( p . 1 9 5 ) . C . O n t h e d i l f e r e n c e si n t h e rnorphology of belemnite rostra(p. 196).Hercalso"alveolar slits"(p. 200).D. On thedevelopment of belemnite rosrra(p. 203).E. The phragmocone andthe sheath(p. 208).F. The pro-ostracum (p. 210).G. Reconstruction of the belemnite shell(p. 211).H. Reconstmction of thebelemnite animal(p.
considered a corroborated scientific insight. The structure of the phragmoconeindeed more readily pointed to the orthocones.The general decapodan characterwas much less obvious.It was Voltz (1836. J a h r b . , p . 1 8 5 ) w h o f i n a l l y r e c o g n i z e di t : " T h e belemnitesare surely so close to the decacera(Blv.)
213).On lif-estylc(p 220).I. Thestratigraphic distribution of (p.221). belemnites K. On belemnite (p. 223).lI. systematics (p. 22a'). Specialaspects
that they should be united with them. They were doubtlessswimming cephalopods,just as the nautilids
I.
p. 176) has been taken as proof of the dibranchiate
were gastropod-like cephalopods". Ever slnce Buckland(1836)the occurrence ofan (195) ink sac (cf.
a.
Preliminaryremarks.
nature of belemnites.Consider,for example,a remark
Belemnoidssimilar to the genusBelemnitesLam. 1801
by H. G. Bronn (1836, Jahrb., p. 40). He also
belong here. The type of the genus should be B.
discoveredfossil ink togetherwith rhyncholitesin shell
paxillosa Lam. (194) 1801 (: B. paxillosus Montfort 1808 : B. paxillostrs Schloth.,in part - B. mtrcronatus
limestone and concludedthat "they probably come
Schloth.1813, 1820 : B. mucronatusBlainv. 1821:
conclusion was partly erroneoussince rhyncholites
Belemnitellamucronatad'Orb. 1845, p. 449, Pl. 33,
belong to the tetrabranchiatesand have nothing to do
Fig. 1-6). This specieshas been established according
with the ink. But in principle the reasoning is
to the descriptionof Belemnites conicus Breyt 1732, which (p. 44, Pl. 8, Fig. 1-7) containsvery good figures
rernarkable(cf. p. 24). ln contrast,Quenstedt(1849) did not acceptthe idea that belemniteshad an ink sac
of Belemnitella mucronata.The speciesthus shouldbe
and therefore had to
calledBelemnitespaxillosus Lam. But we would prefer
cephalopods",i.e. dibranchiates. He could not see the
to conserve the commonly used name Belemnitella
wood for the trees; his exceptional knowledge of
d'Orb., althoughwe list the family accordingto the
details did not permit hirn to see the general c o n n e c t i o n(sc.i . p . I 7 3t .
legitimate designation,again for the sake of tradition. Belemnites(henceforthabbreviatedas B.) is here taken as a collectivename for all Belemnitidae,the generic
f r o r n n a k e d a n i m a l s ( i . e . d i b r a n c h i a t e s ) " .T h i s
be considered "naked
116 b. Diagnosis.
c. On the differencesofthe outer form of
Belemnitidae are moderately slender,often somewhat
belemnite rostra.
pro-ostracum, stocky belemnoidswith a tongLte-shaped
Let us first look at the most frequently observed
the rostrum showing a concentrically layered and
fragmentof belemnites,the rostrum.Its generaloutline
radially fibrous structure, formed by a regular
can be very different among known forms: the earliest
alternationof dense,thin lamellae and transversely-
form is a shorlcone(8. acutus).Inthe youngertypesit
striated (prismatic) intermediate lamellae (cf.
may become long and slender (8. acuarius) and
Coeloteuthis o. 111).
cylindrical in the middle part (8. puzosi), with a spindle- to club-shapedend (.8. clavatus) and grade into laterally compressed,sometimes exotic forms
We first give a generaloverview of the fossil uraterial, before going into detail in order to sketch,stepby step, a life-like picture. In the great majority of casesonly the rostrum of
(.Duvalia).Thereare ail possibleintermediateforms. The biological significance of these large differencesshould not be overestimated.Looking at Figures 61, ll
and J2, one will recognizethe rostrum
the extinct belemnitespecieshas been preserved.More
as a skeletal part of the relatively minor terminal
rarely we find parts of the sheath with the conical
process! which we interpret as a protective and
hollow ("alveolus") in which the phragmoconewas
balancing device (p. 192). Its special shapeis (197)
(cf. p. 175). The most posteriorparts of the iocatedT3
clearly less important than its mass in reiation to the
chamberedshell may be preservedinside this hollow,
p h r a g m o c o n e ,o n w h i c h a h o r i z o n t a l s w i m m i n g
whereasthe more anterior ones were generallybroken
position depends(loc. cit.). The samecan be said about
off or had been dissolved.In exceptionalcasesintact
the secondarytexture of the outer surface.The cracks,
phragmocones are preservedmore or less closely
grooves,longitudinal lines, imprints of vessels,slits, a
united with the rostrum, and in a few specimenseven
delicate point or a blunt end, all these can only have
the (196) pro-ostracumis preservedtogetherwith these
very limited importancefor the living animal. Even
parts (Fig. 87). More frequentlyone finds displaced
taken by themselvesthey should not be given great
phragmoconesof large species,but they always lack
importancein systematics. From the palaeontological
the delicate posterior end. In particularly favourable
point of view, e.g. a distinction of families or
conditionsone may find the pro-ostracumtogetherwith
subfamilies on the basis of the sornewhatvariable
isolatedchamberedshells,either as an impressionor
courseof weak, often hardly visible longitudinal lines,
with the shell preserved(cf. Figs 63f and 65a). The
rs not (198) acceptable.Despitehis very careful study,
correlation of such fragments with the rostrum is
E. Stolley(1919,p. 5l) shouldnot be followed in this
difficult to prove, however, and hypotheses in this
matter. In contrast, his analytical overview of the
respecttend to be unwarranted(cf. p. 171). The best
different sculptureson the surfacesof belemniterostra
demonstration of the three main parts preserved
are very useful. How great the need indeed was for
together is observablein B. brughierl Miller from the
such an overview is shown by the whole literature. I
Lower Lias y (Fig. 66) and Bel. puzosid'Orb. from the
thereforefollow the route indicated.
Oxford Clay (Fig. 87). In the former the whole anitral
1. The apical furro--. Whereassome of the oldest
is fairly distinct in outline, in the latter only the shell is
belemnitesfrom the Lias show smoothtips, circular in
preserved,and both can be reasonablywell completed
cross section,most of the younger ones show more or
from observed fragments. These two fossils are
less distinct longitudinal furrows, which die out
particularlyuseful for our specialreconstructions,since
anteriorly. One can distinguishin particularthe widely
they can be further generalizedon the basis of our knowledgeof shell structure(p. 168).Sinceboth types
occurring dorso-lateraland ventral apical furrows (Fig. 85). (Dorsal. ventro-lateraland numerousintermediate
of background have already been used for the
fuirows also occur in certain species,but they are of
reconstructionof the belemnoid type (p. 167), the
lesserimportance).Often these featuresare limited to
resultingrepresentationis indirectly basedon the same
the posterior end. But they can be much longer and
facts.
may secondarily reach the alveolus, so that the distinction from other types of furrows (see under 2
lt7 Fig. 69.
Schematicillustrations of thc morphology of
belemniterostra. l-10. lateralviews (seenfiorn the right side), 11-16. cross sectionsbehind the alveolus, l7-19. crosssectionsof the alveolarpart,20-25.crossscctionsofapex. ,/. straightshoft cone(8. densPhi1l.{cf. Coeloteuthi.s i 1. 1a. slightly inflated short cone (8. engeli Werner lcf. Nannobelus]). 2. short cylindrical cone (.8. brevilbrmi.s Voltz lcf. Brachvbeltrs,\'1. J. long cone (8. tripartitus sulcatus Quenst. {cf. Salpingoteutltis)). 4. rod-like cone (8.
crcuariu.s gracilis Quenst. {cf.
Solpingoteuthis l'). 5. pole shape(.8.poxillostrsSchloth.\cf. Passaloteuthis|). 6. rod shape(.8.porrec'tusPhill. {ct. C,vlindroteuthisl). 7. blnnt club shape(.B.clavatusSchloth.{cf. Hostitesl). 7a.pointedclub shape(8. host.ttusBlainv. {cf. Hibolites}). E. finger shape(8. irregularis Schloth.\cf. Dactyloteuthisl). 9. slightly roundcd long cone shape (8. gig. ventricosus Querrst.i cl-.Mcgorettrhi s i 1. 10. shorl coneshape(8. contpressusStahl 1cf.Pleurohelu.sl). -11.circular (.8.acutusMiller .lcf.Nannobelusl). 12. compressed(8. c'onpre.s.sus Stahl lcl Pleurobelus,1). 1 3 . c o m p r e s s e dr e c t a n g u l a r( 8 . e x i l i s d ' O r b .
lcf.
Rhabdobelus|). 14. compressedin two parts (8. bipartitus Blainv. {cf. Pseudobelus\). 15. sub-quadratic(8. zieteniWcrner \cf . Broch.vbelusl). 16. flat-bellied(8. venn'oplanusVoltz lcf. Gastrobelusl). -17.deepeneddouble lateral furrows (8. nitidus Phill. {cf. Cvlindroreuthisl).
t
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18. dorsal alveolar furrow (-B. conophorus
Onn
!cf
Conobelus)). 19. ventral alveolarfurrow with adjoining slit and dorsalkcel of conotheca(8. mucronahr.s Schloth.l,cf.Belentnitellal). 20. roundedapex(8. acutus.c'/ovatu.s'). 21. dorso-lateraland ventral apical furrows (8. tripartitus sulcatusQuenst.)(as in 3) 22. with an additional ventro-lateral apical furrow (8. q uin qtrestrIca ttrsBlainv. ! cf . Megat e uthisl,). 23. with additional dorsal and acccssoryapical lurrows (.8. giganteuscrassLts Wemer, ibid.). 2 4 . a p e x c o m p r e s s e dw . ith a dorsal keel (Acroteuthis apicicarinata Stolley). 25. ventral apical furrow (8. puzosi d'Orb.
{cf.
Cvlindroteuthisl).
and 3) may virtually disappear.(c1.Fig. 89a-c).
dorsally shifted "lateral furrows" attain a width of 3-4
The significanceof apical furrows for the life of the
mm and show a complex structure:the furrow proper
animals must have been minimal. I presume that the
extendsanteriorly far beyond the alveolus, in shifting
tegumentalenvelopeof the rostrum was rather tough,
dorsally, whereas it becomes uneven behind the
perhapsreinforced by tendon-like bands. These parts
alveolusand soon expires.Its upperlimit showsup as a
may have had insertionsin the apical furrows. The fact
fine double groovel the lower limit is simple and very
that completelybroken rostra (cf. Duval-Jouve1842,
shallow, accompanied by a low (199) longttudinal
Pl. 10, and our Fig. 82) did not fall off but were
ridge. The whole is an elongatearea with a shallow
repairedcertainlysuggestsa strongenvelope.
depressionin the middle; morphologicallyit can be
2. The double lateral grooves.Much more constant and widely occurring were the generally shallow
interpretedas follows: tight bands [tendons?],perhaps parlly muscular,extendedlaterally along the transition
furrows which occupied the greaterpart of the lateral
from the alveolus to the rostrum; they left these
flanks of elongaterostra,and which were double as far
furows due to the constantstretchingof the bandsand
as well preservedspecimenscan tell us. We can best
resultinginhibition of growth. Thesefurrows disappear
study them where they are most distinct. namely in the gentrsBelemnitella (Fig. 70). Here the somewhat
anteriorly where the fins may have been inserted, as suggestedby comparativestudiesof recentdecapods
lt8 (Figs 62 and 67). The fin insertion (p. 34) was
extendto the area of the alveolus;I believe,indeed,
originally sited on the shell sac, with which it was
that this is the origin of the actual alveolarfurrows (cf.
articulated by a longitudinal, cartilaginous sliding
Fig. 89). In the Cylindroteuthinaeespeciallyone finds
s u r f a c e . T h i s c a r t i l a g i n o u sb a n d c a n b e m o v e d
Belentnopsis-like rostra in which the ventral furrow
anteriorlyand posteriorlyby integumental muscles.We
extendsfrom the apex to the alveolus,where it finally
thereforeinterpretthe lateral furrows as the imprints of
becomesshaliowand soondies out (cf. Fig. 88). In the
thesemuscles,which gradedposteriorlyinto tendon-
Belemnopsinae the furrou,' becomes increasingly
like bandsand anteriorlyinserledon the fin cartilage.
concentratedon the alveolar part. apparently causing
Often the double lateral furrows become very
the special features of the rostrum which are
shallow or only indirectly recognizable:there may be a
characteristicof the subfarnily (Figs 70, 89). The most
n.rinimalflattening of the surface, a dull or glossy
typical feature of the true alveolar furrowsTa,which in
longitudinal striation,which can only be recognizedin
general do not reach the apex itseli is their close
oblique light or after wetting the surface.ln rostra with
relation to alveolar slits, rather than a mere positional
clear evidenceof corrosionthesetracesdisappear,so
relationship. Here we therefore must review these
that no features remain which reflect the tvoical
structureswhich arepart of the inner rostnrmstmcture.
structureof the soft pans.
5. So-called"alveolars/irs". In the Belemnopsinae
3. The r,esselimprints.In certainbelernniterostra,
(q. v.) the alveolar furrows, which forrn sharp grooves
especiallyin Belentnitellamuc'ronataand Actinontax
along the alveolus, are connected with somewhat
(q. v.) rather irregular. bifurcating grooves radiate
problematic structuresthat are characteristicof this
posteriorly, upwards and downwards liom the lateral
subfamily: they are revealedby splitting [the rostrurr] longitudinallyalong the medianplane,which is easily
furrows; they are probably the impressionsof fblood] vessels, and on the ventral side they form a fine
achieved.An alveolarslit is peculiarin that its inner
reticulatepattern.Close inspectionrevealsthat they do
s u r i a c e i s s m o o t h , s o m e t i m e sg l o s s y b e t w e e n t h e
not in general coincide with the furrows. The trunk
alveolusand the alveolar furrow (in contrastto what is
vesselsfrom which the ramified vesselsderivecleariy
observedin other forms, where the splitting surfaceis
did not coincide r'vith the furrows, but they
coarse).(201) The smooth surfacehas a well dellned
accornpaniedthem.
We know of a similar pattern
outline,the shapeof which is characteristic for genera
frorn the fin basesof recentdecapods.They are always
and species.Sometimesa calcareouslayer covers this
a c c o m p a n i e dd o r s a l l y a n d v e n t r a l l y b y v e s s e l s (especiallyveins) that ramify - in a mannersimilar to
surfacein one half; it is presentbefore splitting. lying inside the intact rostrum. It has been interpretedas a
the pattern on the rostrum observed here (they have only a rudirnentary rostrum or none at all) - on the
rostral layers are indeed interruptedat this level, as if
shell sac and in the muscular mantle (200) on which
delicately cut; they are perfectly in phase with one
p. the displacedfin cartilageslie (seealsoBelopteridae
another(cf. Fig. 89f1).
s4-s7).
speciallamella of the ostracum.We only know that the
6 . T , v p i c a l c o r u o s i o np a t t e r n s . I n a n u m b e r o f
4. The median alveolar fttrrov,s. These lurrows originatein the areaof the alveolusand radiatemore or
belemnitesthe rostrum and sheathhad weakly calcified
less far posteriorly; they are different from the other
animal. Peculiarstructuresmay thus result,obscuring
imprints so far mentioned.We distinguish a "rnid-
the original condition.The best known exampleis from
dorsal alveoiar furrow" the occurrenceof which is
the genusActinocama,r (Fig. 92). Here the alveolar
parts that were readily destroyedafter the death of the
largelylimited to the "Dilatati", i.e. our Duvaliinae(q.
sheath(anterior to the rostrlrm proper) (202) was nearly
v.), from a very broad mid-ventral one ("ventral
always destroyed before fossilization. As a result,
canal").The latter is most distinctnear the end of the
e n l a r g e d " p s e u d o - a l v e o l i "o f t y p i c a l s h a p e w e r e
phragmocone,slowly dying out anteriorly and
formed, e.g. with a rectangularcross section in ,1.
posteriorly.This feature was used by d'Orbigny to
qtradratus.Or the outermostand most posteriorparts of
characterizehis group "Gastrocoeli";it is typical of the
the sheathwere eliminatedand the anteriorpart of the
entiresubfamilyBelemnopsinae(q. v.).
rostrum corroded,so that the site ofthe protoconchsits on a typically conical elevation (A. verus). Similar
As observedearlier, the ventral apical iurrows can
119 Fig. 70,
Rostra of different Belemnopsinaeto
I
,,1
illustrate the general morphology of the belemnite rostmm. a-d. Belemnitella mucronata fior-n the Cretaceous near Liineburg,u,ith rcconstructedphragmoconc..7. dorsalview, 1l.ventralvicw. r'. lateralview Note the prominent apex" and the numerous impressionsof vessels.those marked 7 and B representingmain vessels.Their relation to the lateral furrows is of some importancc, the latter form shallow, elongateddepressions.1. ventral limiting ridge, 2. ventral furrow, 3. very shallow middle furrow, 4 and J. double dorsal lurrow, 6. terminal part of the lateral line proper with vessels crossing irregularly, 7 and B. main vessel fumows. 9. ventralfurrow n'ith alveolarslit. cl. On a median sectionthe alveolarslit (.s)is visible to its full extent. It can be seento extcnd to the phragmocone.But in reality it does not reach the conotheca(c), since it remains separatedfrom it by a differentiationofthe sheathwhich is visible in the crosssectiond1. (Looked at from the inside ofthe alveolus it appearsas a fine double line, which seemsto mark the position of the siphuncle;but it can be easily removed.and one then finds the underlyinglongitudinalridge of sheathmaterial). e.
A correspondinglongitudinalscctionof -8. ha,status. Here the slit (.s)extendsfar posteriorly,cutting through the rostral lamellaeas
ln
d and in Fig. 891. The posteriorend is alwaysindistinct.
structuresare observed in the anterior parts of other
are more specialized: delicatekeels,remainingcloseto
belemniterostra(e.g.Neohiholitesewaldi).
the apex or extendingfrom there towards the alveolus,
Other types of rostra show less regular corrosion
have been observed several times, as have been fine
patternsin their anteriorpart. (cf. e.g. Quenstedt1849,
longitudinal grooves starting at the apex, giving the
p. 444). This is observed,in particular, in younger
falseimpressionof accessoryapicalfurrows.
Hibolites,e. g. sub/itsi/bmls Rasp.
The surface of well preservedrostra in general is
The alveolusis often completelylost; this prompted
smooth and glossy, sometimessuggestingthe presence
e.g. Blainville to assumeits natural absencein certain
of a cuticula.But quite often this conditionis destroyed
s p e c i e s( h e n c eP s e u d o b e l u s q , . v.). In other cases
by chemicalor physical agencieswhich actedbefore or
deepenedpseudoalveoliare formed, since the material
after burial, so that the belemniteshave a dull aspect
surroundingthe posterior part of the phragmoconeis
even if no visible damageis observed.Such damage
less solid. This can be observed in
some
can be typical of certainspecies(203) andmay reveala
C y l i n d r o t e u t h i n a e( F i g . 7 l e ) . T h i s c o n d i t i o n i s
lessersolidity of certain parts or a particular chemical
particularly conspicuouswhen the juvenile rostrut'n,i.e. the axis, is very solid and - after disappearance of the
composition,or else a peculiarity of occurrenceand of the enclosing sediment. Some belemnites appear
surrounding material -
naturallyto have a granularor slightly wrinkled surface
projects freely into the
(Stolley,1911, p. 186,Fig. 1). pseudoalveolus
(cf. Huxley 1864on B. elongatus).
A c c o r d i n gt o S t o l l e y( 1 9 1 9 ,p . 2 2 ) s p e c i m e nosf B . araris Dum. from northern Germany show regular,
d. On the developmentof the belemnite rostrum.
medio-dorsal,longitudinalfissureswhich may extend to the alveolus:they are probably due to the destruction
Quenstedt(1849) alreadyknew that the the shell layers of a belemniterostrum do not follow regularly one
of shell lamellae which then leave irregular gaping
after the other startingfrom the inside (Fig. 71). On the
edges(vaguelyreminiscenlof Dtt'alict).
axis behind the protoconch he found the "small
Other strttctureson the surtaceof belemniterostra
belemnite", which he took for the iuvenile fon.n
t20
OIJNil
Po,@
t':
'l
n-'-t:
{\.
.---::- ..=
{'
{ *_\_-._= _
v, L
[1|xF=:" ek
Lt.
Fig. 71.
Structureand developmentof belemniterostla.
a. Homctloteuthisspinota (Quenst.)fiom the Middle Jurassicnear Aa1en.Original specimenin the BavarianStateCollections(public collections), drawn in natural sizc. Median section. The innennost layers of thc rostrum are not necessarilyvcry precise. so the juvenile rostrummay appearsomewhattoo slcndcr. b. Pachyteuthisabbreviara(Miller) af'terPhillips (1870, Pl. 35, Fig. 92). Arca of the apical line. There the lamellac arc indistinct. suggestinga specialconsistencyofthc "axial thread"(a.r).Greatly enlarged. c. Pas.saloteutlrls sp. ("8. tripartitus Schloth.") after d'Orbigny (Pal. fr. jur. P1.5, Fig. 8). Upper Lias. Slightly corrcctcd median section.Some individual rostral lamellaeare ernphasized;in the alveolusthe growth lines of the conothecaarc visible anteriorto the preservedscpta (se). The conothecais dralr'n as if viewed obliquely from below, so that the width of thc mcdian plate of the proOne gro*1h line (l) is emphasised,so that the outline of the juvenile shell, including the juvenile roshum, ostracumcan be assessed. is recognizable.Thc accuracyis questionable.(205,)
=
t2l d. Brachybelusgingensis(Oppel)after Phillips (1864,P1.5, Fig. 11). Median sectionofthe siphuncle,slightly corrected. e. The same,for B. vulgaris (Y. and B.) after Phillips,p. 22. f. B. insculptus(Phill.) after Phil1ips,p. 46. g. Megatetrthisgigantea or quinquesulcata(Hartm.). Combinedfrom severalspecimens(cf. Quenstedt1843,Pl. 28, Fig. 7). Note the Coelotetrrhis-andNannobelus-like form of the juvenilc rostrum. Middle Jurassic(Dogger cr). l. Typical transversesectionofa belemnitebehindthe protoconch. l. The same,in thc areaof the alveolus.Note the distinctionof the porcellanouslayer (hatched)and the nacreouslayer (white) in thc pro-ostracum. k. Hibolite's hastatus (Blainv.) from the Upper Jurassicp of Treuchtlingen.Original in Munich. The juvenile rostmm (Stolley's "cmbryonicrostrum") especiallyemphasised. L Oxyteuthisspec.Drawn in similar fashion.After Stolley (1911, Pl. 9, Fig. 2). Slightly corected by addition of the natural growth lines. Thc juvenile rostrum only reachesthe protoconch to embraceit in cup-like fashion. (?) It probably should be imagined to continueinto the phragmoconesheath,but this can hardly be checkedgiven its texture. m. Pachytetrthis(Acroteuthis)apicicorinato Stolley, from the Lower Neocomian near Braunschweig,with a similar, but shorter, rather indistinctjuvenile rostrum (r). A later stage(2), by contrast,is very distinct in all individuals and thereforemarks a second, naturalphaseof development. n. P. (A.) oehlmannensisSto1ley,from the upper Middle Neocomiannear Braunschweig.Here the secondphaseof developmentis indistinct. The parts drawn with dotted lines in the vicinity of the protoconch are maceratedand show that the solid part of the juvenile rostrumis indeedlimited to the protoconch(as Stolleyprobablyassumed). o. Reconstructionof the animal for the juvenile rostrum shown in c. It is clear that this stagecould not be an embryo; it was a young animal,perhapsat the end ofthe favourableseasonofthe year ofhatching. The fins can be imaginedaccordingto Fig. 67. p. Median sectionat the phragmoconeend of llibolites hastatusBlainv., after d'Orbigny (Pal. fr. jur., Pl. 19, Fig. 61.About 5,, nat. size. The true primordial rostrum (pr) is visible at the posterior end of thc protoconch"it continuesinto the axial thread (a-r), and foms a typical componentof all the rostrastudied. q. The complete shell at hatching from the egg capsule (3/1nat. size); reconstructionbased on the structure of the tvpical shell nucleus. r. The siplruncleof Hibolites hastatusafter d'Orbigny (ibid. Fig. 7). Calcareousand chitinouscones[i.e. septalnecksand connecting rings] are distinguishable, as in Nautilus.The chitinousconesare stippled. s. Median sectionofBrachybelusgingensisafter Phillips (1364,Pl. 5, Fig. l l). r. Odontobelusbrevirosn'i.s (d'Orb.) after Quenstedt(1858,Pl. 41. Fig. 22), from Boll. Lias !. Comparethejuvenile rosrraln g. u. Dactyloteurhis(?) enigmaticus d'Orb. From d'Orbigny (Pal. fi. jur., Pl. 22, Fig. 1). A late, problematic form of the Passaloteuthinae from Oxfordianmarl (boundarywith the Argovian). (cf. p. 238; perhapsbelongingto Bracht;teuthis). si. siphuncle;co. conotheca;at. protoconch;r/. rostral lamellae (growth lines); ap. apical line; a"r. axial thtead; po. pro-ostracum; h"v. Iateral plate of pro-ostracum ("hyperbolar zone"); r. growth linc of conotheca;.te. septum of phragmocone(the more anterior septa are omitted in the figure, to show the juvenile shell three-dimensionally);jzr. juvenile rostrum;pr. primordial rosffum. Seethe preliminary note on the conditionsdescribedhere (Naef, 1922.Eclogae). v. Split (median?)specimenof "Belentnites"obhrsusBlainv., after d'Orbigny ( 1886,Pal. univ., Pl. 7j , Pal. dtr.Pl. 37, Fig. 10).ci Diploconidae.
("embryo") without studying it in detail or trying to define it. Later observersrecognizedthe continuation
having a well developed, calcified, small rostrum (sepiids) show only a knob-like rudiment of the
of the "small belemnite" as the beginning of the "axial
prospectivespine.In belemnitehatchlingsit may have
thread" of the later parts and sometimescalled the
been more fully developed,but a long, pointed process
whole structure an "embryonic thread" (cf. Stolley 1919,p. 8). It is always seenon well preservedrostra
in this position at an embryonic stageis inconceivable, becauseit would have causedimmediatehatchingfrom
ground down to an exact median plane (Fig. 71p) and
the egg caseTs. The preparationsat my disposaldid not
must be interpreted as the first rudiment. We do not
allow me to see how the "primordial rostrum" grades
know, however, whether it was formed during or after
into the sheath,but I supposeit correspondsto the
the embryonicphase.Hatchlingsof recent decapods
typical condition (Fig. 72). Given the delicate early
r22 parts of the shell, thin sectionswould be necessaryfor
cannotclaim suchbelemniterostrameasuring5-6 cm
a detailedstudy. At any rate, one has to assumethat at
in length as pafi of a (207) decapodembryo. I presume
the time when the roughly conical primordial rostrum
instead that the end of the first year has left that
is formed a certain number of chambers already
indication.Surfaceswimmerslike belernnites(p. 192)
existed,and the sheathcan no longer be limited to the
cannot have been independentof seasonalchanges.I
protoconch.The primordial rostrum is continuedby the
found a similar break in growth (Fig. 7lm) in some
apical line (206), whtch must be regardedas a real elementof the rostrum,not just a mere line76;this is
large forms (e.9. Acroteuthis apicicarinqta Stolley).
corroboratedby some median sectionsand by split
juvenile rostrum from the rest; in the posteriorparl the
specimens.In any event,the rostrum grows apically,
separationis rather inconspicuousor even invisible,
and in the true belemnites the rnaterial added to the
and in the anterior part, i.e. in the area of the
apex apparently maintains its special character
protoconch,which is surroundedas by a cup, the sharp
throughout growth. ln contrast,the main mass of the
boundary also disappears.In contrastto what Stolley's
rostrum shows the well-known. rather consistent
figures (Fig. 7ll) suggest,the rostral lamellaedid not
structure: conical lamellae of
conchiohn are
stop at the protoconch. They must have graded
sequentiallydepositedone on top ol the other, ending
(typically) into the sheath,which may have been very
apically in the apical line. In transversesection they
thin. This continuationcannotbe seendirectly, neither
appear like the annual rings of a tree, but they must
which he in Stolley's figures nor in his preparations,
have been formed at much shorter intervals,similar to
kindly made availableto me for study.I fully subscribe
what we observe in recent cuttlefish. The space
t o h i s p o i n t o f v i e w ( 1 9 1 9 ) r e j e c t i n gt h e u s e o f t h e
between the individual lamellae is always filled with
juvenile rostrum as a special distinctive feature, as
shell material showing a radial structure,at least in the
p r o p o s e db y O . A b e l ( 1 9 1 6 ) . I n t h e r o s t r a l f o r m
rostrum proper. Towards the alveolar sheaththe shell
describedby Stolley, a separatejuvenile rostrun.rdoes
lamellae become more closely spaced and finally
not occur, neither in B. clavat.s nor in any of the
becomeindistinct (exceptin thin sections),whereasthe
relatedbelemnitesfrom the Lias. It merely representsa
fibrous structureremains conspicuous,especiallyon
check in the growth of young belemnites,the
broken surfaces.The sheathappearsto be particularly
distinctnessof which may have been slightly
hard and solid. The surface of the inner layers of the
exaggeratedby Stol1ey. In many well preserved
rostrum,i.e. the juvenile rostra,is first smoothand only
specimensof different species,it is inconspicuousor
later shows the specific charactersof the respective
barely evenrecognizable.
subgroups(grooves,slits etc). The first occur-rence of
One should not expecta very sharpdemarcationof the
A more general interest of juvenile rostra is
the latter is not easily found, since cross sections
undeniableif they differ in shape from the definitive
intersect different lamellae at different levels of
ones,evenifthe changeis very gradual. A b e l ( 1 9 1 6 ) t r i e d t o s e p a r a t et w o b e l e m n i t e
formation. It is easyto discover,however,that grow-thwas not
"families" on the basis of "embryonicdevelopment";
really continuous.In belemnites from the Lias
this is not possible,as stronglyemphasized by Stolley
(Passaloteuthinae) some individual lamellae already
(1919). For Stolley's Polyteuthidae
differ in thickness,whereasthis phenomenonbecomes
Rhopalobelus
much more regular in the belemnitesof the Upper
"Conirostridae", u'hereasthe other groups! which are
Jurassic and
Cretaceous (Belemnopsinae,
are equivalent to
except Abel's
united in Abel's Clavirostridae,could have somecloser
Cylindroteuthinae)(Fig. 71k, i). In particular, the
relationship.A closerlook revealsthe existenceof all
anterior part of a slenderaxial element is boundedby
the gradualchangesfrom conical through cylindrical to
one or severalthicker lamellae;this can be interpreted 'Juvenile rostrum" closing the secondperiod of as the
club-shapedjuvenile rostra; they may be characteristic
growth. This interestingstructurewas demonstratedby
define more fundamentaldiscriminations,which must
E. Stolley (1911) who called it the "embryonic
be consideredartificial. Only the younger belemnite
rostrum". That definition is not acceptablegiven the
groups (p. 2a2 and onward) show an "embryonic
generalconditions of development(cf. p. 103). We
rostrum" sensz Stolley (its ostensiblyspecialcharacter
of individual groups,but they (208) cannotbe usedto
1 a )
tzJ
being regardedas attenuated);the oider types from the Lias lack such a structure.A definitive judgement
1'
i: 9
l' Iia. i \ \ l-v
about the systematicsignif,rcanceof this feature is not yet possible. Some circumstances suggest that the rostrum attained a definite size and mature form. The largest specimensfrom a given locality are mostly smooth, glossy (as if polished), the younger ones are dull. Huxley (1864) finds a cuticula with a wrinkled surface in a fully-grown B. elongatus. e. Phragmoconeand sheath. Earlier authors,including Quenstedtand Voltz, had effoneousideas about the size of the phragmoconeand the continuationof the periostracuurover it. Thus Voitz ( 1 8 3 0 ) d i s t i n g u i s h e st h e C r a s s i m a r g i n a t ai n d t h e Tenuimarginati.In the former the short alveolusis said to terminatein a thick rim, in the latter to thin out on the cone. In fact only the latter situation is real, as far as our evidencegoes.The periostracumlayersdecrease in number and continue, as a thin but solid sheathof the phragmocone,to the free edge of the cone and probably also cover the pro-ostracum.The rostrum lamellae, which are separatedby fibrous layers, become increasinglycrowded and thin anteriorly, but they neverthelesscontribute effectively to increasing the weight of the shell (p. 192). There are no completealveoli sensa Quenstedtand
Fig, 72, Reconstructingbelemnite shells. Development as deducedfrom the structureof the rostrum or alveolusand the phragmocone(Voltz, 1830). a. Bel. rluinquesulcattrs Blainville, an idealizedrepresentatiotl of a common fossil. The figure is partly based on the
others.Perhapsonly the rostrum (in the extendedsense
descriptionby Phillips (1868, Pl. 24); it shows the shearh
given p. 176) is fully preserved,in contrastto the more
with the anteriorpart of the rostrum split open,thus exposing
frequent caseswhere it is broken off anteriorly. We
the growth lines or lamellae. The phragmocone is totally
have alreadyexplained(p. 176) that no remains of the
surroundedby the conotheca,on which the growth lines can
alveolar sheathcan be found on isolatedphragmocones
also be traced. One growth line is artificially emphasized;it
which lack the rostrum. The rostra of belemnitesare rather blunt compared with thoseof the aulacoceratids. As far as I know from m y o b s e r v a t i o n sm a d e o n c o l l e c t i o n s , a n d o n specimens(209) I collectedmyself (see also Werner
is the basis of the reconstmctionof the juvenile shell shown in b. in the latter, the muscularmantle has been addedto the free margin of the she1l,thus completing the mantle sac, the whole being coveredwith the skin. A picture of the complete juvenile animal was achieved by adding a typical decapod headand funnel.As to the fins, seeFig. 67.
1912),their apicalanglesin profile rangefrom l0-30',
c. The supposedshell of a hatchling, with a freshly formed
as a general rule 76-27". However, individual species
primordial rostrum(p. 203).
show greatvariation,whereasceftain generakeep close
d. The embryonic shell prior to the lbrmation of the first
to their normal mean. The best method is to measure
septum,a stagestill observablein teuthoids(Fig. 61a).
the angle after grinding a well preservedrostrum to the median plane of the alveolus.Well preserved,large phragmoconesmay also yield reliable results.Crushed onesare ofno use in this respect. The structureof the siphuncleis not invariable either; indeed it shows large variations which can not
e. A median section through the shell nucleus, showing the juvenile shel1 representedin c. The initial caecum of the siphuncleand the prosiphon are drawn in dotted lines. since they are never preserved. They are assumed to be as in Spirttla(Fig.26).
t24 Fig. 73. - Reconstructingbelemniteshells a. Conothecaof a phragmoconeofB. "giganteus", unfolded on to a single plane to show the growth lines (anterior part), the lateral lines (posteriorpart) and the main longitudinal 1ines.By uniting the edges (10) around an oval core (c) one obtains a three-dimensional reconstruction (disregarding the slight curvatureofthe cone).The figure was obtaincdby fixing a piece ofpaper to the surfaceofa well-preservedpiece ofphragmocone (the original specimenof Quenstedtfrom the Middle Jurassic6) and making a contact copy of the lines. The suture lines in the posteriorpafi (9) were obtainedin the sameway fiom a differcnt specimenin the GeologicalInstituteof Jena;they are lessdensely spaced(but not very distinct) in Quenstedt'sspecimen.Only in the anterior part are the growth lines drawn in roughly natural spacing.In the hyperbolarzone(4) this is not t-easible. b. A contact copy made at the mid-ventral line showing a slight siphuncularsinus of the suturesand the tangentialpoints of the
L 1
septal necks. The points marked (x) correspondwith a and c; they lie in an enlargedinterscptalspace. c. Phraqgmoconcin apical view, to show the distribution of the
"main lines" on the crosssection.1. mid-dorsalline; 2. right half of medianplate; 3. medianasymptote;4 right lateralplate; 5. lateralasymptote;6. lateralarcuatezonc; 7.lateraltangentialline, "lateral line"; B. ventralwall ofphragmocone. d. Reconstructionofthe posteriorpart ofthe body, with the addition ofa rostrum (r) to the emphasizedjuvenileshell (brevis stage) and of a muscularmantle (lrr) to the free shell margin. The shell and muscularmantle are coveredby the skin. Skin and sheathare omitted on the right side of the body so that the rostrum is shown in median section,and the conothecain side view. The apical angle ofthe phragmoconeis 22o.This probablyrepresentsMegateuthisrhenan(Oppel)(p. 2a0).
so far be clearly interpreted according to their
shown in Figure 90 could be confirmed,our knowledge
systematic occurrence (Fig. 11). In the forms
would be greatly increased.The preservedpro-ostraca
resemblingBrachl,belusandMegateuthis,it looks like
are very delicatesheetsof calcified shell substance,the
a string of pearls; in the Belernnopsinaeit rs more
median plate with its feather-like sculpture (Fig. 87)
markedly stretched,similar to the siphuncle of the
being thinner than the lateral plates. Thus they look
Aulacoceratidae.But the septaare more closely spaced
(Fig. 41a-c very similar to the shellsof prototeuthoids
than in the latter (Fig. 95); the lengthsof the chambers
and 42b),to which doubtlessthey arehomologous.
are one fourth to one tenth of their widths. The
Nowhere can the whole free shell margin be
conothecashowsgrowth lines and delicatelongitudinal
observeddirectly (as in Acanthoteuthisspeciosa').And
lines, which in fact are linear elevationsthat are most
that would be necessaryfor a completereconstruction
distinctin the areaofthe lateralolates.
of the animal. This reconstructionmust of course dependon a previous reconstructionof the shell, based
(210)
on the detailedkno'vledgeof its parts. f. The pro-ostracum.
Correspondingto the typical, spindle-shaped mantle
Little direct evidenceis availablefor the pro-ostracum
sac, the pro-ostracummust have been slightly curved
of belemnites.Our Figures63, 87 and 90 containabout
along its longitudinal axis. As it graded into the
all that can be shown on the basis of well preserved
conothecaduring development,it must have caused
rnaterial. A striking feature is the similarity of the
ventral curvature of the end of the phragmocone;this
isolatedpro-ostracumfrom the Lias (Fig. 63b) to the
can indeedbe observedin all belemnites,though to a
pro-ostracumstill united with the phragmoconefrom
variable extent. The curvature(at least initially) never
the Upper Jurassic(Fig. 87).
attains a degree comparable to that observed in
If the hypothesis underlying the reconstruction
sepioids;thus it doesnot becomesecondarilyinhibited
r2s by the developmentof the rostrum (cL p. 47). The
on the hypostracum and even better on the
latter is perf'ectlycapableof maintainingthe straight
periostracum,namely on the inside of the alveolus
growth of the apex by compensatorygrowth on the
(Figs 63c and 71c). When looking at isolated
ventral side. The eccentricgrowth of course has to
phragmoconesof large species,one rnay obtain a
change in accordancewith the curvature; this can
particularly clear picture by tracing the growth lines
indeedbe observed(cL Fig. 71). This resultsin a dorsal curvatlu'eof the apical line that counteractsthe ventral
directly on to paper wrapped tightly around the cone (Fig. 73). They can then be reconstructedin three
curvatureof the cone.diminishinggradually.
dimensionsby bending the paper correspondingly;the slight curvatureof the cone of course gets lost in this
(2tr)
way. Likewise the secondariiyaddedmaterial(p. 210)
g. Reconstructionof the entire belemniteshell.
forming a pattern (feather-likestriation) on the pro-
T r u l y c o m p l e t eb e l e m n i t e s h e l l s h a v e n e v e r b e e n fbund. However, a fairly accuratereconstructioncan
ostracumis not visible in such a tracing. Only the juvenile phasesof the pro-ostracumare retainedon the
now be achieved on the basis of the evidence
conotheca.Thereforeobservationsmade on isolated
rnentioned a b o v e( p . 1 6 8 - 1 7 1 )i.n d e e dm o r e p r e c i s e l y
fragments,and those from the cone (Fig. 631) have to
r.towthan u''ashitherto possible (compare our Figures
be integratedinto a reconstruction.lt is especially
7 1 . 1 2 a n d7 3 w i t h t h o s eo f e . g .Z i n e l 1 8 8 5 ,p . 4 9 8 ) .
important to note that, (213) due to the density and
The result is fully corroboratedby some relatively
steepcourseof the hyperbolic lines, the relative length
completefossils(Fig. 87). The rarity of suchspecrmens
of the structurecannot be easily calculated;it is often
is not surprising.A glanceat the whole shell showsthat
assumedto be shorterthan it really is. Thus Phillips
its preservationintact is highly unlikely. After leven natural)deathof the animal the shell must have drifted
( i 8 6 3 , p . l 7 a n d 1 8 ) p u b l i s h e du n b e l i e v a b l pe i c t u r e s that have been reproducedmore than once (Fischer
at the sea surf-ace,thus being exposedto wave action
1 8 8 7p, . 3 6 2 )( c f .F i g s6 3 ,7 l - 1 3 , 9 0 ) .
and to being destroyedespeciallyby (212) surf. Only detached rostra and pro-ostraca had a chance of
h. Reconstructionof the belemnite animal.
becoming rapidly buried. Thus the most posteriorpart
Since the belemnite shell can be reconstructedrather
of tl.rebroken phragmoconewas likely to sink with the
precisely, on the basis of careful interpretationof
rostrum; larger parts of the phragmoconeprovided
combined fragments,we have a solid basis for the
sufficient buoyancy to the rostrum to float as long as
reconstructionol the animal itseif. Following our
the chamberswere not pr.rnctured.
specialconditions(p.22), we can add the mantlesacto
We have seen (p. 177')a number of belemnoid
the shell, and for the other parts we can proceed
fossils lacking the rostrum; in most casesthe other
according to our Figure 62, modifying it whenever
parts of the sheathare also missing. It thus appears
necessaryon the basisof evidencefrom specific fossils
likely that they representedthe corpses of widely
(Figs 66, 67). We inevitably end up with the same
distributedbelemnitesin which the rostmm was broken
representationas those obtainedfor typical belemnoids
off. This loss may have occurred at death or
in general, and for the "genus Acanthoteuthis"in
subsequentlywhen the dead animal was carried to an
particular (p. 177-184),finding tangible differences
intensivesurf zone. The numerousrostra with clean.
only in the shapeof the rostrum and in the degreeof
intact, empty alveoli (without any remains of the
elongationof the body: basedon what we concludedp.
conotheca)and the more rarely occurring isolated
210, belemniteswith a curved,excentricapical line and
phragmocones(from which the sheathin generalwas
curved phragmoconemust have had a strongly arched
r''ery cleanly separatedby maceration)show that a perforatedor broken sheatheasilybecameisolatedT?.
back, i.e. a less extendedform than others. Moreover
What is not naturally available can be extracted
we can deduce from the large apical angle of the phragmoconethat theseanimalswere clearly stocky, in
fi'orn the specimensby reading the growth lines of the
the extreme case corresponding to the condition
phragrnocone. as shownby Voltz (p. 168).Theselines
mentionedabove (Passaloteuthinae; cf. Fig. 72). The
are particularly distinct on the outsideol the ostracum,
slender,especiallythe club-shapedrostra are associated
but if the latteris lost they also shou'up as impressions
with phragmoconeanglesof less than 20o, and they
126 reconstructions,general proportions are not given
t.i{,,h,&l
precisely, although they can be obtained from the fossils,and there is no systematicdescriptionof typical forms or correlation with soft parts, which can be achievedon the basis of the rnorphology of recent forms. It is not acceptablesimply to combinea vaguely assumed shell form
with
an equally vague
r e p r e s e n t a t i o no f a c e p h a l o p o d . T h e s c h e m a t i c r e p r e s e n t a t i o ngsi v e n b y d ' O r b i g n y ( 1 8 4 0 ) , O w e n ( 1 8 4 2 ) , Q u e n s t e d t( 1 8 4 9 ) , H u x l e y ( 1 8 6 4 ) , P h i l l i p s ( 1 8 6 5 ) ,O . F r a a s( 1 8 6 6 ) ,P o h l i g ( 1 9 0 9 ) ,a n d E . F r a a s (1910)cannotbe acceptedas scientificreconstructions (cf. Figs 14-16). Comparedto them the newer attempts offer some progress(Fig. 80) since they reflect an effort to improve knowledge of the recent decapods and to use them as a basis for comparisonwith fossils. N e v e r t h e l e s sa s p e c i a l e r n p h a s i so n p r e c i s i o n i s wanting, as is the capacityto systematicallyrecognize the typical features within the diversity of recent forms78.We indeedhad to acquirethe prerequisitesfor Fig. 74, - A reconstructionof the belemniteanimal after d'Orbigny 1840(1eli)and 1842 (right).Here one recognrzes that author's understandingthat belemnites*-ere typical d e c a p o d s .S p e c i a l ' m o d e l s ' f o r t h e p i c t u r e s w e r e I l l e x coindeti andAlloteuthis subulata (Lam.)(p.217). The proostracumcorrespondsto a prototcuthoid shell (.Leptoteuthi,s?
(215) suchan emphasisby many years of study; it was only thus that we created the basis lor potential success. by the following steps: Our methodis characterized 1.Study the shells as completelyas possibleand reconstructthem on the basis of the seneral laws of grorvth(p. 168).
cf. p. 120)(from Abel 1916,p. 219).
2. Take preservedremainsof soft parts (Fig. 66) into account, considering that ink sac, show minimal curvature and excentricity. Their
muscularmantle, impressionof the head with
reconstruction reveals extremely slender animals
traces of mandibles.arms with hooks must
(similar to Fig. 71o, but even more markedly elongate).
exist, although not necessarilyproviding a
(cf. Figs 72b and67c,but also 95).
complete picture, in general conformity with
Under these circumstancesother belemnite species
typical decapodorganisation.
can be easily reconstructed,without yielding new
3.Use this type (Fig. 62), which is well defined,
insights. We therefore are content with this general
for a methodical comparisonon the basis of
presentatron.
the diversity of form of decapoddibranchiates
We neverthelesswish to add a few critical remarks about historical descriptions,becauseIhey serve(214) to illustrate our intentions.
whenever comDiementarv information is needed(p. 7).
Reconstructionsof
4. Consider that the special adaptationof the
belemniteshave been made in large numbers; they
typical organisationto become a more
form part of the standingfurniture of palaeozoological
specializedshell form can only be discovered
and palaeontological lecturesand textbooks.None of
through circumstantial evidence about the
them can claim nrore than merely historical interest,
structure, environmental conditions and life
althoughsome of the most recentones(by Stromervon
style of similar recent animals; (216) rn
R e i c h e n b a c h l 9 0 9a n d A b e l 1 9 1 6 , s e e F i g . 8 0 )
addition to illustrating a natural life fom one
correspondgro,s,tomodo to the presentviewpoint
has to take account of secondaryfeaturesof
except for the number of arms. But even in these
structureand behaviour.which asain must be
=
127
Fig.75. - Other old reconstructions of belemniteanimals(fiom Abel 1916.p.22 I ancl223). Flon.rleii to lighr accoldingto the f o l l o w i n ga u t h o r sa: . O w e n ( 1 8 4 3 ) ;D . Q u e n s t e d(t1 8 4 9 ) ;c . H u x l e y ( 1 8 6 4 ) ;d P h i l l i p s( 1 E 6 5 ) . A s o l i d b a s i so 1 ' r l o r p h o l o g i c a l knor"'lcdgeis not recognizablein thesefigures (only Huxley's figure shows a relatively natural l'ierv). lNote: 'fhc figure shor.i'sonll three reconstructions. c was in fact copiedby Abel from Zittcl (1884);Huxley (1864) did not incluclca lcconstrlrction.Phillips' ( I 865)versionwas copicdby Abel ( 19 16, p. 221, Fig. 90) but not by Nael.
seenin a harmoniousrelationshipof parts within
ecologicalterms,the former examplc(rvhichcouid be
a form determinedbv certainlawsre.
imaginedto beara relativelyheavybelernniteshell:cf. p . 1 6 5 ) ( 2 1 8 ) a p p e a r sm o r c c l o s e l , vs i r n i l a r t o
Figure 77 shows two Mediterraneandecapodsthat are
b e l e m n o i d s . H o u ' e ' , ' e r .s u c h s p e c i a l s i n i i l a r i t i e s
reminiscentof the outlinesof belemnoids.so that an
concerningcertainpartsof the body shouldnot be used
idea of belemnitesin life may be supported:a) is a "nektonic" species,a good s$.immerfrom the group of
uncriticallyfor the reconstructiou of unknou,n(extinct) whole animals(cf. p. 7). becausethis r.r'ouldcarry tlie
loliginid squids.The strong elongationof the posterior
risk of projecting even insignificant peculialitiesof
part. however, is a feature of mature males only and
l i v i n g f o r r n s i n t o t e n t a t i v e r e c o n s t r u c t i o n s .T h e
has to be interpretedlike other external features(217)
mechanicaiand ecological signilicanceof the parts
of sexualdimorphism(apparentlydevoid of essential
consideredshould be scrutinizedrnore caretully.and
functions in reproduction).Since this feature is absent
the possibility (219) of combined functions of such
in young males and in females,and also often less
p a r t s s h o u l d b e e x a m i n e d f i r s t , a p r e r e q u i s i t et o
marked,it cannotbe regardedas an individuallyvital
"palaeobiology"sensu O. Abel. But even lrore
organ8O. Once present, it naturally contributes to
important is a methodicalelucidationof typical
stabilisationand steering.b) is a planktonicjuvenile
relationsas explainedearlier(p. 8); a palaeobioiogical
oegopsidsquid.The elongationof the conusand ol the
treatment can only be attempted as a secolldary
neck aid buoyancyin this extremcly delicateanimal. In
a m p l i l i c a t i o n . T h e l a t t e r s L r g g e s t st h i s : t h c
rnorphologicalterms (consideringthe conus) it is rnore
Chiroteuthidaewith their specialjuvenile form (Fig.
closelysimilar to the belemnoids;in physiologicaland
77c) show a typical oegopsidconus(cfl Gonatus.Fig.
t28
ar
tn tr
vd, ad po
Plt
v
a,
b.
Fig. 76. - Relatively recent reconstructionsof belemnite animals (from Abel 1916. p. 226). From 1eft to right: a. Aftcr E. Fraas (1910).b. After Pohlig (1909).c. After v. Stromer(1909). The last one reflectsan undeniableeffort to makeuse ofbasic insights into comparativeanatomy.The number of arms (6), and the lack of knowledgeof the insertionof the muscularmantle into the shell (p. 22) inevitably make the picture rather deficient. tD. ink sac; tt1. tentacle; tr. funnel; po. pro-ostracum;ph. phragmoconelro. rostrum;km. 9111; sl. siphuncle:.ll shell fold; rd. foregut;/rl. mandible;ar. at1-t1s.
59c) in an extremely, atypically elongate form,
in general,including their ontogeny.The latter cannot
although the observed relics of chambers can be
be used, however, by palaeontologistswho lack
consideredtypical. It is doubtful whether these relics can be derived directly from an ancestralteuthoidform
morphologicaltraining (Naefl 1921, On structureand life style) - apartfrom the problernof eroneous data in
(p. 159); at any rate the chiroteuthidsshow a secondary
the literature. For example, the 6-armed brachial
resemblance to belemniteswhich does not permit any
c o m p l e xo f y o u n g o e g o p s i d(sp . 1 6 0 ,F i g . 6 1 ) d o e sn o t
s p e c i a l c o n c l u s i o n st o b e d r a w n , c o n s i d e r i n g i n
teach us anything about their ancestors;it cannot be
particular the delicate, gelatinous nature of these
taken as a phylogeneticinheritance.Otherwrseone
planktonic-nektonic animals living in rather deep
would have to assumehominid ancestorswho at first
water. Abel made much of Chirothaumamacrosoma
had only two teeth,then four, etc. endingup with 3281.
and of the related Ch. imperator (Figs 78 and 80c).
o20)
Adopting Crick's (1902, 1907)assumptionsabout the
As to the lifstyle of belemnites,seeabove(p. 191)
number of arms did not improve the results.We have
on belemnoidsin general.Here the questionwhether
r e j e c t e dt h e m e a r l i e r ( 1 9 1 1 , 1 9 2 1 , S y s t e mp . 5 2 9 ;
the rostrum effectively counterbalancedthe buoyancy
Cephalopoda vol. I, p. 133)and now (cf. p. 27 and 182)
o f t h e p h r a g m o c o n ei s o f m a j o r i m p o r t a n c e .T h e
must reject them even more decisively.That Abel's
(superficial)answer,often provided, insistedon heavy
picturesare often better than those of his predecessors
rostra which would exclude a life style of active
(except Stromer's) is due to his, partly successful,
swimming. The palaeontologists arguing thus (p. 20) seemedto have no idea of the (221) relative size of the
attemptto understandthe body forms of dibranchiates
t29 phragmocone(Fig. 90). The calculationsgiven by Abel
I
(1916,p. 166)at leastshow the oppositesituation;only
I
in speciallyadaptedforms can horizontal swimming be achievedbelow the surface of the water (since, in contrastto what Abel thought, the phragmoconecould not be partly filled with u'ater;cf. p. 11).However,in addition to the rostrum, the considerableweight of the conotheca.the septaand the siphunclehave to be taken into account. The specific gravity of the rostrum and probably of the other shell parts as well is close to 2.675 (Fischer 1887, p. 362), like that of other rrolluscanshe11s.
i. The stratigraphicdistribution of belemnites. The geologicaldistributionof belemnitesin time and spaceis far lrom being well known. The following statements,at least, can be made safely: the true belemnitesappearin the Lower Lias (upper cr) and occur up to the Eocene.There are no reliable finds either earlier or later. They are mainly known from Europe,but they occur in all partsof the world in strata of the sar.r.re age;until we know more about the species .nve involved cannot achieve a precise picture of their phylogeny. They are most widely distributedin the Upper Lias of Germany and England. As to their first o c c u r r e n c el,e t u s l i s t e n t o o l d Q u e n s t e d (t 1 8 4 9 , p . 393). the rnan who knew the Swabian Jurassic in greaterdetail than any other worker of his time: "The belemnites first appear rarely in the Arietetid limestonesof Lias cr, and
with a few interruptions
attain their greatestdevelopmentfiom the Numismalis mar1s82 up to the Jurensisbedsr.Here the number of fragments is immense,indeed there are few creatures that stand comparisonin this respect;consideringthat each fragmentwas surroundedby a considerablemass
Fig. 76. - Relatively recent reconstructionsof belemnite
of flesh, one can imagine that at the boundarybetween
animals(fiom Abel 1916.p. 226). From left to right: a. After
Lias s fl-ower Toarcian] and ! fUpper Toarcian], w h e n e v e r y t h i n g E aw a s d e p o s i t e d i n a c a l n . r
(1909).
envlronment,enormousquantitiesof flesh were slowly brought up by the sea. After this event their number
E. Fraas(1910).e. After Pohlig (1909).c'.After v. Stromer The last one reflects an undeniablecffort to make
use of basic insights into comparativeanatomy.The number of arms (6), and the lack of knowledgc of the insertionof the muscular mantle into the shell (p. 22) inevitably make the
suddenlydecreases; in the Upper Brown Jura fMiddle
pictureratherdeficient.lb. ink sac;tr. tentaclc;tr. funnel;po.
Jurassic] they once again increase in numbers, to
pro-ostracum;p/2. phragmoconc;ro. rostrum;km. glll: si.
linally becomeincreasinglyrareEl."
siphuncle;sl shell fold; vd tbregut;ki. mandible;.7r.anns.
(223) The descriptionsof repairedrostra illustrated in Figure 82 are of generalinterest.They show that thesestructures(p. 199) had a tough covering,or else
They also argue in favour of a littoral life style; under
the
such conditions v i o l e n t c o l l i s i o n s s u c h a s t h o s e
fragments
u.ould
have
been
lost.
130 Fig. 78.
Chirothaunta maclosotnaafter Goodrich (1896)
lrom Abel (1916. p. 173).
This is a chirotcuthidfrom
East Asia, with a slightly swollen posterior end which extends beyond the fins. This end does not contain a rostrum; it enclosesthc older, posterior part of the conus and bears small skin folds ("accessory fins") on the surface,a secondarystructureobservedin ceftain members of this family. There is no reason to assume(rl'ith Abel) their presencein belemnites(cf. Fig. 80c); thcse are very peculiar, rarely occurring structures.which have been a r b i t r a r i ley n l a r g e idn t h i sp i c t u r e .
Fig. 79.
Longitudinal sectionof the
"gas chambers" in the gladius of Chirothauma imperator Chun (after I
C h u n , 1 9 1 0 ,P l . 4 1 , F i g . 1 3 ) .E . e n d o f the visceral sac. S. scptum. Z. "gas chamber".
The chamber formation is
real in chiroteuthids; it correspondsto that observedin gonatids(p. 157). Chun docs not say anything about air included in the charnbers,so this may be a (perhapsfbrtunate) addition by Abel.
documentedhere can be more easily imagined.They also show the vital energy of these animals. (In captivity cuttlefishsoon die when the posteriorend is damaged,whereas under natural conditions they often survive after serious damaqeto the shell and soft body).
e
k. On the systematicsof the belemnites. We have seenthat the very diverse belemniterostra have a relatively uniform basic structure,suggesting that the animals which secretedthem are ciosely related.The unquestionablyconsiderabledifferences are not overwhelming when consideredin terms of their biologicalsignificance.The greatestbiological differencelies in elongationwhere it relatesto a shift of equilibrium or a change of body shape. Differencesin transversesectionappearto be almost ofno consequence.
131
Fig.80.
Belemnitereconstructions in Abcl (1916.p.225. Figs 96-98).The picturcsshow the follovningspccies(fiorn lcfi to right): a. An adult Salpingnteuthi.sacuaria (Schloth.)from thc Lias e. D. The same spcciesprior to.rostrurnelongatron.c.lJibolires semihostattrs(Blainv.) from the Upper Doggcl(!) of S*'abia. d. Hontaloteuthisspinata (Quenst.)fiom the Lorver Dogger-([j) of Swabia.Note that a is a modilledAlloteuthissubulatd(Lam.)(cf.p.2I1): D is the colresponding juvenile stageic is a Cltirothuuno nlacrosonla Goodrich (cf. p. 218); d is an ommatostrcphid,perhapslllex coincleti(Vdrany) u'ith an incolrcctly reduccdnumber of arms to suggesta supposedbelemnitecharacter.This way of reconstructir-rg specicsis unacceptable,evell though relativcly latural forms areproduced.
We will therefore lollow an old tradition and
nomenclature,becauseof the need to integratethe
consider the belemnites as one single farnily of
group in a wider context,ll'hich is deterrninedat f-amily
belemnoids, trying to arrange the speciesv,ithin this
l e v e l . S o m e o f S t o l l e y ' sl a r n i l i e sm a y t h u s b e c o r n e
family. For many aspectswe will rely largely on the
subfamiliesand even lou'er level groups.ivliich is a
careful studiesof virtually Lrnlimitedmaterial by M.
purely formal issue.
L i s s a j o u (s1 9 1 5 ) ,W e r n e r( 1 9 1 5 ) E , . S t o l l e y( 1 9 1 9 )a n d
A different question is whether the systetnatic
v. Billow (1920). If we fbrmally modify Stolley's
features so carefully defined by Stolley pennit a
system in a few points, it is fbr basic reasonsof
simplificationof the system.I indeedthink that this is
132 Fig. 81, -Specimen of a "belemnite battlefield" showing Hastites t'lavalus fiom the Lias e ILower Toarcian]of Bartenbach,Swabia. Slightly reducedin size. Original specimenin thc PalaeontologyInstitute of the University of Vienna. After Abel (1916, p. 20a).
This specimen
may representeither a mass strandingof belemniteshells or the regurgitatcd remains of belemnites from the stomach of an ichthyosaurianwhere such massesof rcmains arc oflen found.
Fig.82.
Regenerated belemniterostra,fiom Abel (1916.p. 215 and 216),afterDuval-Jouve(1842).Thc thickcstspccimcnbelongs
to Duvalia lata (Blv.), the remainingonestoHibolites subfusiformls(Rasp.);naturalsize.
r33 the case, and I thereforeunite e.g. the families
GenusOdontobelusn. g. QryramidalisZiet.')
Pachyteuthidae, Cylindroteuthidaeand Oxyteuthidae.
Gerus MegateuthisBayle (gigantea Schloth.)
(224) The relevant characters(lateral furrows) are not sufficiently important in rnorphological or biological terms to warrant distinction at higher
GenusBrachybelusn. g. (breviformlsVoltz) GenusHomaloteuthisStolley(.spinataQuenst.) Subfamily4. Cylindroteuthinae nov.
systematiclevel; it is rnore irnportantto emphasize
GenusCylindrotetrthisBayle Qtuzosld'Orb.)
natural connectionsthan to insist on subtledifferences.
Genus Pachyteuthis Bayle (excentralis Young
For detailsolthe argumentseethe following section. The systernaticcontributionsof earlier and more recent authorswill appearin the review. Incidentally,
andB.) G e n u s O : r , y t e u t h i sS t o l l e y ( b r u n s v i c e n s i s v . Stromb.)
this survey expressesthe effect of, and correctionsto,
GenusAtrlacoteuthis Stolley (absol ut iformis
an earlier system which first subdivided the old "genlrs" Belemnites (much like Ammonites) into
GenusRaphibelttsn. g. (aciculaMiinst.)
"sections",then genera,and finally into "families". Our
Sinzow.) S u b f a m i l y5 . B e l e m n o p s i n a ne ov.
approach does not of course provide a definitive
GenusBelemnopsisBayle (bessinad'Orb.)
system.The whole material needsa stringent analysis
GenusHiboliles Mayer (hastatusBlainv.)
based on internal and external charactersEs; only such
GenusB elemnoconus (.baudouini d'Orb.)
an analysiscan determineconnectionsand limits and
Genus Parahibolires
flnd the right place for each"species".The significance
Stolley (duvaliaeformis
Stolley)
of the old, still frequentlyused "sections"in relation to
GenusMesohi boI it es Stolley (minaret Rasp.)
our partly new groups will be given in the relevant
Genus1y'eolr ibolites Stolley (semicanaliculatus
chapters.
Blainv.) G enusB eIemni teIIa d' Orb. (.mtrcronata Schloth.) GenusActinocamaxMiller (verursMiller) GenusDicoellles Biihm (meyrati Ooster)
ll. Detailedarrangementof the Belemnitidae.
Subfamily6. Duvaliinae(Pavlow as family) emend. GenusDuy a I i a Bayle (l ata Blainv .)
Contents:Systematic overview(below).a) The sublamily H a s t i t i n a e( p . 2 2 5 ) . b ) C o e l o t e u t h i n a( pe. 2 2 9 ) . c ) (p. 230).d) Cylindroteuthinae Passaloteuthinae (p. 242).e) B e l e m n o p s i n a( pc. 2 4 7 ) . f ) D u v a l i i n a e( p . 2 5 7 ) . g ) (p. 259).Review(p.260). Bayanotcuthinae Systematicoverviewwith norrinal types
GennsPsetrdodtnal i a Qtolygonalis Blainv.) G enusPseudobeItts Blainv . (bip art i ttts B lainv.) GenusConobelasStolley(conophomsZttt.) Subfamily7. Bayanoteuthinae nov. Genus Ba,vanoteuthis Mun.-Ch. (rugifer Schloenb.) GenusS/yac oteuth is Crick.
Subfamily L Hastitinaenov. Genus-IlasllresMayer (clavatusSchloth.) GenusRhabdohelus n. g. (exilisd'Orb.) Subfamily2. Coeloter.rthlnae nov. GenusCoeloteuthisLrssalous(ca|car Phill.) Subfamily3. Passaloteuthinae nov.
a. The subfamily Hastitinae nov. Whereasalmost all known Lower Jurassicbelernnite rostra can easily be related to B. acutus Miller (the oldest form in the EuropeanLias o.: Sinemurian),a new type appearssuddenlyin Lias l3 (Pliensbachian)in
GenusNarnobellr.s Pavlow(acutusMiller)
the form of B. clavahtsSchloth.,simultaneouslywith a
GenusPassaloteuth i.\ Liss. ( brughierl d' Orb.)
more abundantoccurrenceof the older type; the origin
(22S)Genus Pseudoha.stite.r n. g. (scaDroslr^s Phill.)
ofthis new type is totally obscureas far as our present
GenusGaslrobelu.sn. g. (retttroplunusYoltz)
knowledge goes86.lntermediate(226) forms must be
GenusPletrrobelusn. g. (totupresstl"iStahl.)
sought in older strataor in other parts of the world. In
GenusSa/plrgoteuthisLiss. (/r'l.v/11lcata Blainv.)
any caseB. clayatusmust be consideredthe type of a
GenusDaclr'loteuth is Bavlc-( illi,.qrlra'l.lSchloth.)
specialsubfamily, not united with the Passaloteuthinae
134 f ig. 83.
On the morphology of thc hastites and of some r/2 nat. size.
l.clemniteswith abnomal growth.
i.l
i,l lil
d. Hastitesclayatusafter Phillips.Pl. 3, Fig. 7 1"', with distinct leteral furrows (.r, r1).:a1 and a2'.cross sections in the alvcolar region. 1t.Conespondinglongitudinalsection.
li
t . H a s t .m i c r o s t y l n(si b i d .P l . 1 3 .F i g . 3 1 ) . " afler Quenst. 1849, Pl. 29, Fig. 4la, tl . Hast. "sub./irsiJbrmis "shows no ventral furrow" and shows all the characteristicfeatures
$i
l{r
of hastites.From the Malm y nearNusplingen.
lt sh l t
e. Hast. clavatus with thick club, al1cr Friren 1868; el: median
ltt
scctionof'alveolarpart, magnified.
jc.
.1.B. irregularls after Bayle (Pl. 28, Fig. 7) with strikingly elongate juvenile rostrum.
$ld
g. B. "pistilliformis" afterd'Orbigny (Pa1.1i. ct cr6t.,Pl. 6, Fig. 4) with secondarythickeningof the club. as is typical for Ha.ytites. h. B. (Neohibolites)ntinintusList. After d'Orbigny (ibid. Pl. 5, Fig. 9) with secondaryrod-shapedelongationof rostrum,as normally observedin the Acuarii (Quenstedt1849,Pl. 24. Fig. 12a).
(seebelow) as suggested by Stolley (1919,p. 12,39).
H. clayatusis explicitly n-rentioned as typical and the
The separationof this group and the persistenceof its
next speciesare essentiallythe same as those which
basic characterthroughoutthe Mesozoic is probably
Stolley already consideredto belong here. We must
due to the wide-ranging value of the club-shaped
therefore accept Hastites as valid fbr the Clavati,
rostrum(cf. p. 192).
despitefurther restrictions.In generalthehastttes(227)
Hastitinae lack apical furrows; at best there rs a
are rather small (Fig. 83) and they rarely occur in great
weak ventral furow. In contrastdouble lateral furrows
numbers(Fig. 81). Despitetheir sirnilarityLoHibolites
are often distinct, sometimesdeep, and there may be
they differ in outline from the speciesofthat genus:the
lateralkeelsin the stemzone.
bulk of the club is situatedcloser to the end, which rapidly becomespointed.The alveolar sheathpart of
The genusHastitesMayer 1883,s. reslr.
the rostrum is strikingly slenderand short, suggesting an extreme compensationfor phragmoconebuoyancy
Belemnitesclosely relatedto the type of B. clavatus
(p. 192) in hastites. ln other elongate rostra further
Schlotheim(1820,Pl. 2, Fig. 32-33)belonghere,ever
adaptationsseem to play a part as well. We therefore
since d'Orbigny 1842 (Mayer-Eymar1883, Zittel
view the hastitesas clearly nektonic forrns of the open
1885,Werner l9l2) they have been groupedtogether
seas;the scarcity of specimensmay be due to this
a s " C l a v a t i " . P a v l o w ( 1 9 1 3 ) i n t r o d u c e dt h e n a m e
mode of life.
Rhobalobeltts,which was adoptedby Stolley (1919).
Related forms are: B. microstylr.sPhillips (cf.
Since Mayer had lumped all sorts of belemnitesunder
above Fig. 83c), B. neumarktenslsOppel see Zittel
"Hastites", justification for the name may appear
1 8 8 5 , p . 5 0 5 , F i g . 6 9 1 ) , B . s u b c l a v a t u sY o l t z , B .
doubtful. We therefore quote from the original text
privatensis Mayer, B. toarcensisOpp., B. pistillfbrmis
(Mayer-EymarI 883,p. 642):
Blainv. It is noteworthythat frorn the beginning(Lias
"The genus Hastites contains, in addition to the
p-y) the long- and thin-stemmedforms occur (Werner,
typical specieswithout a ventral canal, the subgenera
Pl. 10,Fig. 13);I do not considertheir directderivation
Hibolites Montf., Duvalia Bayle andBelemnitella
from Nannobelus,e.g. via B. charmouthensisMayer
d'Orb. with the two form seriesof H. clavatus". He
andB. alveolalzs Werner, as probable.Such extremely
cites as examples:clavatus Schloth.,charmouthensis
"clavate" rostra placed at the very posterior end of the
M a y e r - E y m ,. m i c r o s t v l a s P h i l l . , t o a r c e n s i s O p p . ,
phragmoconerequire a special structure (228) and
neumarktenslsOpp., bi/er Mayer-Eym.,subclavatus
strengtheningto be biologically functional,and such an
Yoltz, royeri d'Orb., souichi d'Orb., /ischeri Eichw.;
adaptationwould have to be recognizablein stepwise
13s transitionsin the zone of Lias $ to ^1.Such transitional
like Duvaliinae(Fig. 93) is striking. A derivationof the
stagesarenot known to date.
younger belemnites,at least of the Belemnopsinae
The problem is of some importance since Abel (1916) attemptedto divide the whole belemnitegroup
from Hastitinae,via such intermediatetypes cannotbe
into two lines, one associatedwith Irlannobelus,the
excluded. For the Cylindroteuthinaethe relationship is less obvious (Fig. 7l l-o!). - lf theserelationships
other with Hastites(Clavirostridae-Conirostridae). Like
could be fully confirmed, this would vindicate the
Stoliey(1919)we considerthe systematicrelationships
general basis of Abel's opinion (wirh important
to be less sirnple; without agreeingon all points with
r e s t r i c t i o n s ,i n t h a t t h e s e p i o i d s , a u l a c o c e r a t i d s .
Stolley's criticisrns (cf. p. 208), we think the justification of such a simple scheme by a
belemnoteuthids,vasseuriidsand the Coeloteuthinae
schematicallyrepresentedembryonic developmentis
have two main groups of Belemnitidaes. str., which
premature.
would be associatedwtth J'ilannobelusand Hastites.
There is no clear information on the phragmocones
would have to be excluded).We then would indeed
respectively,and the questionto be answeredwould
and alveoli ofhastites, which are rarely preserved.The
only be whether or not these types had a common
remainswhich I have seenhad an apical angle of less
[email protected]).
than 20o. whereasin all the older belemniteswe find angles ranging 23-30o, most often 26-27". (The
b. The subfamilyCoeloteuthinaenov.
y o u n g e r g r o u p s B e l e m n o p s i n a e ,D u v a l i i n a e , a n d
Here we place a number of peculiar rostra flroln the
Cylindriteuthinaealso have anglesbelow 20o in their
Lower and Middle Lias of England, Gennany and
typical representatives; only Pachyteuthis is more like
France (Fig. 84i-n). Whereasthe Hastitinaeshow an
the Passaloteuthinae in this resoect).
early climax of developmentof the sheath.we see the weakestexpressionof this developmentin the present
The genusRhubdobelusnov. gen.
group: the Coeloteuthinaetotally lack an elongated rostrum; the latter appearsin its simplest fbrm as a
Here belong the forms relatedto B. exilis d'Orb. in the
thickened end of the sheath.similar in a way to the
uppermostLias and lowermostDogger.According to
assumedancestrallbrm of the belemnoids(p. 166);we
Werner (.1912,p. 115) they shouldbe associated with
will find it again in the phragrnoteuthids(?) and in the
Hastites.In fact the speciesmentionedis thought to be
(Fig. 67, p. 186).Were it not for the belemnoteuthids
"gradually derived" from a variety of H. clavatus.If so
typical radial structurereminiscentof the belernnites
it would have to show a long, rod- to club-shaped juvenile rostrum, something that has not yet been
related to Nannohelts, the forms under discnssion
observedin the adult form. Werner (loc. cit.) regardsB.
parlicularly striking that the typical concentriclayering
paryus Hartmann as a juvenile form; while Stolley (1919,p. 34) finds that thereare sufficientnumbersof
of the sheathand the apical line are said to be lacking
"very slenderand thin juvenile forms" of undeniableB.
q. v.). relationsIo Belemnoteuthis,
would have to be excluded fron-rthe family. It is
(Fig. Sai). (Completely lacking? Abor-rtconceivable
exilis. At any rate, Rh. parvus andserpulatzrsQuenst. (1885,Pl. 41, Fig. 19 and 20) mustherebe excludedas
The genusCoeloteuthisLissajous1912.
independentspecies. The typical forrr is rod-shaped,the cross section
At present we have only one genus, Coeloteuthis
quadrangularanterioriy, whereas the more or less
L i s s a j o u s( 1 9 1 5 , p . 1 3 ) , i n t h i s s u b f a m i l y ; i t u . a s
markedly thicker posterior end has a rounded cross
o r i g i n a l l y c r e a t e db y L i s s a j o u s( 1 9 1 2 , p . 9 . ; a s a
section. The stem bears lateral lurrows of variable
subgenusof B. calcar Phillips (Fig. 841,m; Lias y of
depth; they lie closer to the dorsal side. Slight ventral
England and France).(230) B. excavatusPhill. from
and dorsal furows may aiso be present.None of these
the samelevel in Englandand Swabia(Fig. 84i, k) is
furrows extendsto the posterior end (Werner loc. cit.,
apparentlyclosely related;perhapsit simply represents
but seeFig. 93c).
an older individual of the samespecies.Both show a
(229) The similarity of thesefbms to P.seudobelus-
sub-quadraticcrosssection.In contrast,the fbrm called
j
136 B. calcar Phill. of Figure 84n is a different type, which
angle of only 18' in both lateral and dorsal views; it
would rather recommend amalgamationwith B. dens
possiblybelongsto a giant form of Hastitinae.
P h i l l . ( 1 8 6 4 ,P I . 2 , F i g . 6 ) . T h e l a t t e ri s a f o r m f r o m Lias 13,which was also known (from Swabia) to W e r n e r ( 1 9 1 2 , P l . 1 0 , F i g . 7 ) . T h e s t r a i g h tc o n i c a l
The genusNannobelusPavlow 1913.
shapeis common to both; the furrowed surfaceof the
Here belong belemnitesrelated to B. acutus Miller
latteris probablydue to post-moftemdamage.
(Fig. 84), i.e. rather small, pointed rostra of short to
Two opinions are possible for the interpretationof
slenderclub shape(cf. Wemer 1912,PL.10, Figs 1 and
this genus: either we have before us the surviving
4), without a columnarpart and without distinct apical
primitive forms of all belemnites(i.e. only slightly
furrows, of limited excentricityand roughly oval cross
modified descendants of the ancestralspecies),or we
section, rer-niniscent of a strongly rounded triangle.
are looking at secondarily simplified relatives of
When the generally narrow dorsal side becomes
Nannobelus.Ineithercaseone might view theseforms
broader the triangle may become quadrangular.The
as the predecessors of Belemnoteuthis.
s p e c i e s a n d v a r i e t i e s h e r e a s s e m b l e dd o u b t l e s s comprise the earliest belemnites.Nannobeltrs acutlts
c) The subfamily Passaloteuthinaenov. 87
occursas early as the Sinemurian(Lias upper o,). There
Here belongbelemnitesallied to B. paxillosusSchloth.
are some records of belemnitesfrom older strata,but
1 8 1 3 , i . e . i t s c l o s e s tr e l a t i v e su n i t e d i n t h e g e n u s
they seem to be erroneous,as shown by M. G. Fabre
Passaloteuthl.s.Son-reare less advancedin that they
( 1 9 0 3 ,B u l l . S o c . G 6 o l . F r a n c e{ 4 } , t o m e 3 , p . 2 a 9 ) .
preservethe form of juvenile rostrum assumedfor
(Seethere earlier communicationsby M. Haug and M.
Nannobelus, others go much further in that the rostra
Kilian, p. 245-249). Very similar forms have been
becomesecondarilyelongated,endingup rod- or even
found in the Lias ft (N. oppeli (Mayer)) and Lias y (M
club-shaped.This group is limited to the Middle to Upper Lias and Lower Dogger.In the Upper Lias (e-6)
armatLts{Dumortier}). A strong breviconic form is N. e n e g e l i W e r n e r ( 1 9 1 2 ,p . 1 8 0 , P l . 1 0 , F i g . 4 ) . T h e
they show an enormousdeployment,both in terms of
apical angle of the phragmoconein lVannobeltrsranges
numbersof individualsand diversity of species.But
from25-2J".
distinctionbetweenthesespeciesis difficult. Perhaps the Passaloteuthinaeform the stem group of all the following subfamilies; at any rate they are a
The genusPassaloteuthlsLissajous1915
particularlyvariableunit. juvenile rostra are particularly The Nannobelers-like
Here belong the belemnitesof the Middle Lias related
characteristicof the diversity of the subfamily. Their
to B. bnrghieri d'Orbigny, which is a well definedtype
degree of elongation (231) ts variable, within limits
among the fonns surroundingB. paxillosus Schloth.
s i m i l a r t o t h e v a r i a t i o ni n N a n n o b e l u s . I n t h e a d u l t
( c f . K e f e r s t e i n1 8 6 6 ,P l . l 3 l , F i g . 8 , Z i t t e l 1 8 8 5 ,p .
rostrawe never find rod- or club-shapednuclei, even
504, Fig. 688 and my Figure 85b, g, h). The genusis
though some adult rostra may show a decreaseof
identicalwith Holcoteatlls Stolley (1919, p. 35) and
relativelength(Fig. 83f).
comprisesthe forms generally united as "Paxillosi"
In this subfamily the phragmoconesare strikingly blunt, in contrastto those of the Hastitinae.The apical
(Deslongchamps 1878,Mayer-Eymar1883,Quenstedt 1885, Werner l9l2). They first appearin the Lias .1
angle in general ranges from 26-2Jo, but it can be as
(Pliensbachian)with a wide distribution;they are close
small as 23" (8. virgatu.s tsee Passaloteuthisl and irregularis {seeDactvloteuthisl accordingto Werner
to Nannobelus. B. alveolatus Werner from the
1912) or as high as 30'(,8.
slenderjuvenile rostrum clearly resemblesN. acutus,
pvramidalis, see
Lotharingianrnust be considereda predecessor. The
Odontobelus). However, I lound a fragment of a
but the amount of elongation is very variable1.(233)
rostrumwith a round, large alveolus(2 cm in diameter)
more or less in the same sense as the slender-
in the Lias e fl-ower Toarcian] near Holzmaden,(232) which was reminiscentof B. paxillosrrs.but has an
cylindricalrostra of the adults(cf. e.g. Phillips 1866, P l . 1 0 ,F i g . 2 6 S : P . l a e v i sS i m p s o na n d P I . 6 , F i g . 1 6
r37 Fig. 84.
Some belemniterostra consideredto be similar to thc
o. /'.'--1
/-"'/:\
prototypeof the Passaloteuthinae.
f--_ .3
KJ!t/
l. Nannobelusacutus(Mlller) from the Lower Lias of Lyme Regis.a:
l1[bffi
crosssection;b: median section;c: lateralview showing lateral furrow
l[Wr
(especiallydistinct in this specimen).c1: 1y'.in/irndibulunt(Phill.) for
".tEllla.l
comparison.Apcx curved, dorsally and vcntrally fincly striated; r/: juvenile lbrm correspondingto a (more slender);e: ventral view ofa,
h$/\ It/ V
afterPhillips(1867,Pl. 1. Figs I and 3). 2. B. brevirosrrlsd'Orb. (Pal. Fr. terr.jur., Pl. 10, Figs 3-6). (This is probably tlrc juvenile form of Megatefihis, cf. Quenst. 1849,p.422);
rut ' & r---a
IJ
l/
/
\ f \t
Y/ n \"fl \l d..
v
lt
ll
n, 1.,'=h
./: median section;g: cross sectionwith siphunclc(egg-shaped);ft:
\
I i
\/
n
(''/ i
dorsal view with dorsolateralfurrows; next to it the cross sestion of the apex.Probablya cmshedspecimen. 3. Coeloreuthisexcovoto (Phill. 1867, Pl. 2) liom the Lowcr Lias of Lyme Regis; i: longitudinalsectionwith (filled) alveolus;probably not exactly median, hence the rounded end of the alveolus. without protoconch; /r: lateral view (from the right side) with shallow lateral furrow;1: cross sectionof apex: ru: a different specimenwith distinct dorso-lateralfurrou's; Phillips thereforeplacedit in a different species (8. calcar). ,1.n: a form tentatively identified as Coelctteutltiscalcar by Phillips (Fig. 5); its regular conical shapcrequiresspecialrecognition(nov.
t. v l
1l
spec.?cf. p. 230).
5. Various forms rcsemblingNctnnobelus,after Quenstedt1849.o: "8. bretis" (Nannobelus.tclttlts)from the Lias cr (Pl. 23, Fig. 17b);p: "8. breviJbrmis"fiom the Dogger e (?) (Pl. 27. Fig.27a); q: B. " t r i p o r t . b r e v i . sf"i o m t h e L i a s e [ L o w e r T o a r c i a n ]( P l . 2 6 . F i g . l 8 ) ; r : " f r y o f t r i p a r t i t u s " f r o m t h c L i a s e f l o w c r T o a r c i a n ] ( P 1 . 2 6 , Fig. 29); s: "8. compressasVoltz" from the Doggera (P1.27,Fig. 10a);t:"8. acutus"juv.; zr:the same,old specimenfrom the Doggera (Pl. 27. Figs I 4 and 17a). Comparewith thejuvenile stagesof B. quinquesulcatus tn Quenst.(Pl. 27, Fig. I 2). p-Lt are hardly identifiable; thcy rnay belong to Oclontobelas (p. 238) or in part, as juvenile forms, to Megotetrthis and r/2nat. Scr/pingoteuthis. Al1 drawings size.v: ventral,d: dorsal.
S: P. apicicurvatd).The excentricityis insignificant,
blunt phragmocone angle also varies strongly, from
the cross section is oval in outline with only slight (231) compressionwhich appearsmore pronounced
about23oto 28o,more generally26-27".
dorsally. In addition to the speciesalreadymentioned, known (althoughnot always well defined) speclesare: P. carinata (Zieten), P. nigra (Lister), P. elongata
The genusPseudohastitesn. gen. Here belong Passaloteuthls-likeforrns which look
(Miller), P. virgata (Mayer), P. milleri (Phi11.),P.
closer to Hastites due to the elongation and club-
/aseola (Dumortier), P. ctpicicurvata (Blainv.). The youngestspeciesis apparentlyP. whitbyensls(Oppel) from the Upper Toarcian.- ProbablyB. trabecula Liss.
shapedthickening of the posterior end. However, a we find there (Fig. 85fl) is not like B. scabrosusPhill.
slight swelling is also observedin Paxillosi.But what
1915alsobelongshere,althoughit showssomeoverall
(Fig. 88e), and we thereforemake this speciesthe type
similarity to Cylindroteuthis. The shapeof the apex is
of a new genus.Unfortunatelyit is a unique specimen,
very different; it may be short, pointed, globular, or
but it is so clearly def,rnedthat no uncertaintyremalns.
elongate, sometimes straight, sometimescurved dorsally,with apical furrows (dorso-lateraland ventral)
The three apical furrows in particularprohibit inclusion in the Hastitinae.- B. charmouthensis(p. 226) may
or without such furrows. A slightly club-shaped
alsobelonghere.Localitiesareprobablyin Lias y.
thickening is often seen, and such variations (though less marked), aiso occur within species.The rather
138
@.
@^@
ffiffi m \n-lb.
i*
li;,iY
\--lr
li:$
1 .S,
\ . . g v' 4/ 1 .
l'la
@..
[d,.
,l^
J'.,-l
@,.
k/.,
ffit
%1,
@ tl
W.
@
ft$
t$
i.u
[\i$L$
Fig. 85. On the formationand modificationof thc paxilloserostrum1r l nat. size). "8. o. brevifbrntis amalthei" - Brach.t:beluszieteni (Mayer) fiom the Lias 6 near Hechingen, which is the typical fbrm of Brachybelus(cf. Fig. 84 o), after Quenstedt1849(Pl. 24.Fig.22). apicicunata (Blv.) from the Lias y, which in the juvenile phaseis similar to Fig. 84a. h. "8. poxitlosis numisntalis" - Pu,ssaloteuthis D1:the samevicwed from the apex. shou'.ingthe typical apical lurrows. 62: a /ounger specimen.rnorerounded(ibid. Pl. 23, Fig. 21a, 2lb.22b\. (Stahl)from the Lias 6 near Heiningen.c2:apicalview (ibid. Pl. 24, Figs 18aand b). c. "8. colnpressus"- Pleurobeluscompressu.i ventroplanus(Yoltz) fiom the Lias 1,,seenfrom the right side.dl: apical view; c/:: crosssectionof d. B. t'entroplanus- Ga,strobeltts middle part (ibid. Pl. 23, Fig. 20a-d). analthei" (same as a) from the Lias 6 near Hechingcn, older specimen.c1: frontal view of alveolar end; e2. e. "8. brevifor-mi,s terminalview of the apex(ibid.P1.24,Fig.21a-c). (Dumortier)ftom the Lias 6 near Gross-Esslingen.li:from apex(ibid. Pl. 24, Fig. 2a. b). f. "8. elongates" Pa,ssaloteuthis.faseola ntilleri (Phill.) from the Lias 6 near Breitenbach(ibid. P1.24, Fig. 4). Reachcsat least g. "8. paxillosus arnalthei" - Pas.soloteuthis twice the size(and triple thickncss)in the Lias of England. poxillosa (Schloth.)from the Middle Lias of England.after Phillips, Pl. 20. Fig. 52. h. "8. poxiltostrs" - Pas,saloteuthis i. "8. tligitalis popillatus" DacO-loteuthisirregularis (Schloth.)fiom the Lias e fl-ower Toarcian] of Heiningen.l1: cross section; 12:lounger specimen(ibid. Pl. 26. Figs 4a.3, 1a); i3:D. iruegttlarlsafter Phillips, Pl. 15. Fig. 37a seenliom the apcx. Lias of England. fr. Samespeciesfrom the samesource(P1.15,Fig. 39), especiallyelongatespecimen. (Stah1)from the Middle Lias of Engiand (after Phillips, Pl. 3, Fig. 8), ventral view. /1: cross section of l. PlettrobehrscompressLts alveolarregion; /2: lateralview from the right; /3: crosssectionof a smallerspecimenliom the SwabianLias 6 (after Quenstedt1849. Pl. 24, Fig. 20c). This form is so reminiscent,in terms of shapeand cross section,of the Duvaliinae(Fig. 93b) that a close relationshipseemslikely. However, this could only be considcredas demonstratedif the furrow were dorsal at /, and if a juvenile rostrumwere visible inside(detailsthat may not havebeenrecognizedby Phillips).
The genusGastrobelusnov. gen.
Posteriorlythe ventral side is slightly curved upwards, without furrows, at best with flat lateral areas. The
Here belongs B. ventroplanu.\Voltz (Fig. 85d) as the type. The pecuiiarity of this form justifies a generic
posterior end can be club-shapedor more evenly
distinction. The blunt posterior end and the strong
cylindrical. The apical line is strongly curved and excentric(2:1). The phragmoconeangle is blunt (235)
flatteningof the ventral side of the rostrum are striking.
(about 26"), the alveolus is circular. This species
t39 occurs widely in Lias upper T and lower 6 of Swabia and Bavaria;it has also beenfound in Alsace and in the
elongation,upon which a once more well organized envelope of layers was added. The basic form is
R h 6 n e b a s i n ( c f . W e r n e r 1 9 1 2 ,p . 1 1 6 - 1 1 7 ,P l . 1 0 ) .
slender-conical.FollowingLissajous(p. 18) we regard
Related varieties or speciesare B. suhdepressus YoTIz
B. trisulcatus Blainv. as the typical species. We
and (?) B. unthilicattisBlainv.
a s s o c i a t ew i t h i t : B .
The juvenile forms are
slender,more cylindrical,lessflattenedand excentric. The position of Gastrobelarsis uncertain.Werner
b r e y i s u l c a t u s Q u e n s t . ,B .
longistrlcattrs Yoltz, B.
tricanaliculatu.s
quadricanaliculatusZieten,B. acuarius macer Quenst.,
unites it with clavatus, i.e. with our Hastitinae,but
B. tripartitusstrlcatusQuenst.,B. unisulcatusBlainv.
there is no good reason for doing so. The internal
( ? ) , B . s u l c v s t i l u s P h i l l i p s , B . t e s s o n id ' O r b . : B .
structurearguesagainstsuchan affiliation.
blainvillei Desh.(Bayle, Pl. 30) may also belonghere (cf. p. 245 and Fig. 89). In any event, u'e have to
The genusPleurobelusnov. gen.
include B. acuarius Schlotheim and allies, namely B. gracilis
Here belongsB. compressusStahl as the type; it is the opposite of the previous group, so to speak. See
(Stahl) Zieten, B. acuarius ventricosLrs
Quenst., B. tubularis Young and B. lagenae.fbrmis Zreten and draw attention(seebelow) to the relation to
Werner (1912, p. 117) who describesthe rostrum as "club-shapedin lateral view. This shapeis due to the
Dact,vlotetttlrls Bayle. Young specimensof Salp.
rrodification of the stem, which still has a nearly
often look
square cross section close to the phragmoconebut posteriorlyis compressedin such a way that the dorsal
irregtrlaris (cf. Bayle). But they can also terminatein a pointed apex, as in Bel. regularr.rPhill. (Fig. 86d), so
side becomes narrower than the ventral side. This
one should not pay too much attention to this
acuaria and fragments lacking the (broken off) apex strikingly
similar to Dach'loteuthis
c o m p r e s s i o ni s t h e c h a r a c t e r i s t i cf e a t u r e " . . . " T h e
similarity. Other sirnilaritiespoint Io Aulacoteuthis.
posteriorend in generalis blunt and facing dorsally. It
Megatetrthi,s,and Odontobelus. In terrns of their
bearstwo short, shallow dorso-lateralfunows, whereas the 'stem' is adorned on either side with two.
ontogeny the group shows a peculiar intermediate position: as far as is known, the juvenile rostra are
sometirnesseveralstreaksextendingto the posterior
tnore slender than in Megateuthis, t'norelike an
part, the two most ventral ones being the best
elongatedNannobelusac:utLts, similar to what we find
developed". The phragmocone angle is 25o.
in speciesof Pas,galoteuthis.An intermediatestage
Occur:rence in the Lias 6 of Swabia,Bavaria, northern
reminiscentof Dact.vloteuthis is not generalized. Abel
Germany,the Rh6nebasin and England,and also in the
(1916) designatedB. acuariu.sas the type of his new
black Alpine limestonenearCorps.
genusCa.rplteuthis.Consideringthe formationof the "tubulus", that speciesshould be included here 1cf.
This form also has an uncertainposition. If the juvenile forms are l{annohelu,s-1ike, as suggestedby
Ztttel 1885,Figs 618,619 and 687, p. 50a).The apical
Werner (Pl. 11, Fig. 6b), it shouldbe united with
angle of the phragmoconein Salpingoteuthisrs 25-21'.
Passaloteuthlsand could be easily relatedto forms like
in Dactvloteuthis itis only 23o(Werner1912).
P. virgata.If not, it shouldbe viewed in relationto the Duvaliinae, although they have a blunt phragmocone
The genusDactyloteuthl'sBayle 1878.
angle (25'). (In the speciesunder discussionit is only 2lo)
Here belong B. irregttlarls Schlotheim(Fig. 85i) and its allies, a group with problematicrelationshipsto the
The genusSalpingoteuthl'sLissajous1915.
precedinggenus.Its demarcationfrorn Brachybelus
A number of species,some of which are problematic,
includesB. regularisPhill. (1886.Pl. 15,Fig. 38). The
belong here; their peculiarity is (236) that the solid, short juvenile (Odontobelzr.sto Dactyloteuthis-like)
blunt end with (237) a small, superimposedapex (cf.
part of the rostrum is foilou'ed by an indistinctly
be considereda genericcharacter;but the main species
layered (perhapsunlayered) part. u'hose formation
cannot be accommodatedin any other genus without
must have caused a rapid (o'u'er-hastv as it were)
disturbing the picture. Perhaps Bayle is right to
and Passaloteuthisis uncertain.The latter probably
Bayle 1878,Pl. 28), which is often brokenoff, cannot
=
r40
I
rl
xl I pt tffit
@ ar
o
cll
[,/
ii;I llt l
-tn\
tr
6 t Y.:-_,/ r\, /,
V
r\li
tq:
Fig. 86. -Morphology of Salpingoteuthis(o-d1andMegateuthis(e-i').g:
'ia,
'/2 others: nat. size.
a. "8. acuariusgrocilis" after Quenstedt1845,PI.25, Fig.4a liom the left side.Lias e fl-ower Toarcian]Holzmaden.d1i crosS section;.72:ventralview ofapcx. b. "8. acuariustubularius" (ibid. Fig. l0a; bt: crosssection:b2:ventralview ofapex. Lias e fl-owcr Toarcian]Ohmden. c. " 8. octrariusnla.'er" (ibid. Fig. 21. Lias ! Ohmden..'r: fragmentwith juvenile rostrum(Fig. 28 ibid.). (11) there d. "8. ocuarius" from thc Upper Lias. Original spccimcnin Munich teachingcollections.After a solid juvenile rostr-Lrm follows a loosc.apparentlyun-layeredcore (12) and then againa nomally stnrctured,hard cofiex (-/3). e. Distal paft of rostrum in B. ellipticus after Phi1lips,P1.21, Fig. 53. Lateral view. e1: destroyedapex,which shouldbe imaginedas an addrtionto this hgure. ./. Proximalpart of e with phragmoconefragment.Angle 23'. From the early Middle Jurassicof England. g. B. gigonteus ventricostrs(Quenst.)from Bayle, Pl. 25, Fig. 2.Early Middle Jurassic.Probablybelongsto M. oalensis VoilIz.tll nat.size. h. B. actraris."fry" after Quenstedt,Pl. 25, Fig. 10. Posidoniashalesnear Ohmden.A sma1lspecies! probably ajuvenile aalensis,in thc collectionsofthe GeologicalInstitute at Jena.11:anteriorview i. B. gigonteusquinquesulc'attts, with alveolus:a2: differentspecimen;sameorigin. i3i croSSsectionof apexwith typical funows (x, y, z).
combine parl of our Salpingoteuthis with B. irregttlaris
Pl. 22, Fig. 1) deservesspecial attention; it is a
(238) (\? t\ S. lagenaeJbrmis,acuaria, ventricosa, tubularis). These relationshipscannot be clarified
particularly short, rounded form from the Oxfordian,
without a careful comparative study of the internal
phragmoconeis strongly curved and rather excentric,
anatomy of all these species.Perhapsone part should
the apical line is shorter than the dorsal radius of the
be included in Dactyloteuthis,
sheath.The speciescan be placedhere only tentatively,
another in
Odontobelus?l The problematicB. enigmaticusd'Orb. (Pal. fr. jur..
with
a
short-conical juvenile rostrum. The
nowhereelse.It can be regardedas a late branch ofthe Passaloteuthinae ratherthan as a pathologicalfonn of a
=
141 r r o d e r nt y p e . S e ep . 2 0 4 a n d F i g . 7 1 u , a s w e l l a s t h e
The genusMegateuthisBayle 1878.
peculiarB. penicillatusin Phillips(1856,Pl. 1, Fig.2). The availabledevelopmentaldata allow us to place Tlre genusOdontobelusn. gen.
close to Odontobelttr a group of sometimesvery slenderand large belemnites,the type of which should
Here we place the belemnites surrounding B.
be B. giganteasSchlotheim.Longitudinal sections(Fig.
p),ramidoli,sZieten (Quenstedt1849, cf. above Fig.
71g) show that the rostra of this specieswent through a
84g). i.e. the normal (not abnormally elongated)
short-conical,progressivelylengtheningCoeloteuthis
conical "Tripartiti" of Werner (1912). They show a
and Odontobelus stage during post-embryonic
,\,!annobelus-likeoutline (which correspondsto the typical juvenile forms of the following genus)and, as a
development,followed by a stage comparablewith a short Pa.ssa/oteuthis. ln the speciesmentioned,such
very characteristic feature, the three apical furrows
stagesalways show a distinct dorso-lateral,and a less
typical ofrather young Passaloteuthis,one ventral and
distinct ventro-lateralapical groove, in addition to
two dorso-lateral.The form variesbetweena very short cone (the above speciesand .8. brevirostris d'Orb., cf.
which dorsal and ventral apical grooves often occur (Fig. 86i). In this phaseseveralspeciesof Megatetnhis
Fig. Saf or a moderately elongate one (8. conoideus
have been identified or describedas B. quinquesulcatu,s
Oppel,in Quenst.1849Pl. 27 Ftg.4) and more slender conesas in B. tripartitus gracilis Quenstedt(loc. cit.,
Blainv.8E. Subsequentlyanother,sometimesthorough lengthening occurs, as a result of which a uniformly
Pl. 26, Fig. 67). An intermediatesituationis observed
stretchedcolumnarcone may be fomed, often marking
in B. ox1,cs11us Ztet. (loc. cit., Pl.26, Figs l9-21). The juvenile rostra are very short-conical,Coeloteuthis-to
the termination; in other forms a long-cylindrical
N a n n o b e l u s - l i k e , a s i n t h e f o l l o w i n g g e n u s .T h e
the quinquesu/catus stage may receive a narrow
transversesectionis always slightly compressed. It is
("meagre")end posteriorly(Fig. 869).
obvious that these forms are closely related to one
elongationoccurs (Bel. ellipticus Miller). Alternatively
In Megateuthls the greater part of the rostrum is
another and that they representa sort of reversion to
ungroovedand has an ellipsoid(laterallycompressed)
rheltlannobelrlstype, especiallywhen one considers,in addition to the figures mentioned,Quenstedt(1849, Pl.
cross section; similar compression affects the phragmocone.The latter has a suitablesize for detailed
2 7 , F i g s 2 a n d 4 , o n t h e b a s i so f P l . 2 0 , F i g s 1 7 , 1 9 - 2 1 ,
observations of the finer structure, therefore it has
P L . 2 5 ,F i g s I a n d 3 , 1 8 , 1 9 ) a n d W e r n e r( 1 9 1 2 ,P l . 1 2 ,
always been used for structural descriptions.Thus
F i g . 4 , P l . 1 3 , F i g . 5 { 8 . t r i p a r t i t u s c r a s s a s } ) .T h i s group occurswidely in the lMiddle andl Upper Lias (e
Voltz (1830) (240) used e.g. "8. compressus"from
and 6) and lowemost Dogger (cr). - The largestforms
683). The structure of the conotheca can also be
belonging here (239)
observedin detail; it is easyto distinguishthe nacreous
are clearly similar to
Gundershofen: B. rhenonusOppel (Zittel 1885,Fig.
Megateuthis, which should probably be associated
and porcellanous layers on fragments even with the
here.This relationshipbecomesconspicuous rvhenone
nakedeye. Theseforms are of specialinterestgiven the
considersthe figures of B. ventralls published by
combinationof very old, simple types of ontogenywith
Phillips (1866,Pl. l7) showing a form combiningthe
ratherhighly developedones(Figs 72 and86).
f e a t u r e st t r i p a r t i t u s
d i s t i n g u i s h e df t o m q u i n q u e s u l c a t u sb y t h e l a c k o f
The most important speciesof Megateuthisare'.M. ovata (Blainy.), M. yentralis (Phill.), M. opalintts
dorso-lateraland the enhancementof the ventral apical
(Quenst.) (after Werner), M. rhenana (Oppel), M.
furrows. The juvenile stagesare at first Coeloteuthislike, then more slenderNannobelus-like.The apical
elliptica (Miller), M. longct(Voltz), M. aalensis(Voltz) : M. ventricosa (Quenst.),M. crassa (Werner) - M.
angle of the phragmoconevaries with the degree of
gigantea (Phill., Bayle). A collective name, especially
elongationof the rostrum (23-30'). The alveolus is
for the speciesfollowing rhenana, ts B. giganteus
stronglyexcentricand curved.
Schloth.
and giganteus. lt is only
Megateuthis is possibly close to Salpingoteuthis\e (cf. Fig. 86). The apparentlysecondaryelongation of
I
the apex which tends to occur (to different extents)in
t42 Fig. 87.
An especiallybeautifulbelemnitc shell from the Oxfbrd Clay of
England; after Mantell 1848 (cf. also 1850),1/2nat. size.
This is a
puzosi (d'Orb.) showing the rostrum (R), the specimenof Cylindroteuthi.s phragmocone(Ph) and the pro-ostracum(Po) in their natural relationships. thus confirming the ideas of Voltz (p. 168). The reinfbrced lateral plates
Plx
are easily distinguishedon either sidc of the delicatemedian plate with its feather-like striations. The mediar.rplate should be imagined as rcconstructedaccording to Fig. 63b. For the rcconstructionof the whole see Fig. 90. Of great importanceis the direct proof of the connection betweenthe three main parts of a belemniteshel1,which in most casescan only be deduced.In the fossil shown Fig. 66c the exact shapeand structure of the pro-ostracumare still missing, its outline and sculpturebeing uncertain.The figure is from Zittel 1885,p. 501; it is also reproducedby Keferstein.I 866.P1.13I .
crassa). 2. The ventro-lateralones may be missing while the ventral ones are particulariy distinct (ventralis,opalintrs {Quenstedt1849,p. 308, Pl. 42, Fig. 13} in the form illustratedby Werner 11912,p. 1 3 3 ,P l . 1 2 , F i g . a ) ) { c f . J a n e n s c h1 9 0 2 ,P l . 1 2 , F i g . 7)). 3. The ventralgroovesmay be missingor be very indistinct(rhenana). The apical angle of the phragmoconevariesgreatly, dependingon the extent of primary elongation.In M. elliptica it is only 20o, in M. aalensis and others it is 27'. The frontal angle is much smaller due to compression(23 and 16o,respectively).This causesa lateral flattening of the mantle sac, (241) and the animal of M. elliptica may thus have been a slender, elegantswimmer,whereasM. aalensiscan be irnagined as a rather stocky animal, like most of the (Figs 67, 72, 13). Passaloteuthinae The genusBrachybelusNaef 1922. Here belong the short, thick, strongly excentric(2:1) belemnitesfrom the Middle Lias to Lower Dogger, .\
from which Homaloteuthl.ican probabiybe derived.
all the speciesis again achievedby a sort of over-rapid
The shorlnessof the rostra makesthem look similar to
growth, often resulting in the suppressionof layering
Nannobeltts,the more columnar form is reminiscentof
and the radially fibrous structure.Longitudinal sections
Passaloteuthls.The differenceis emphasizedby the
show this to variableextents.D'Orbigny (Pal. fr. jur.
excentricityand the relatedcurvatureof the apical line
Pl. 14, Fig. 1) illustratessuch a form, in which the
(Fig. 71f and s) which often causesa dorsailyangled
elongatedterminalpart is unlayered(markedblack).
apex. The cross section in general is very slightly
The apical grooves vary greatly. Ventral, dorsolateral and ventro-lateral ones may be l
c o m p r e s s e d ,o v a l t o r o u n d e d f o u r - s i d e d . A p i c a l
simply
grooves may be very distinct ventrally and dorso-
complete,or accompaniedby additionalgrooves,e.g. a
laterally. The juvenile rostra vary between
dorsal one (quinquestrlcata,aalensis,elliptica, longa,
Coelotetrthis-and short Nannobeltts-likefoms similar
t43 to those we find in Megateutlris. They tend to be
evidence either of the existenceor absenceof
shorterthan in speciesof Passalotezltlrls.Lissajous
intermediateforms; however,the excentricity(Fig. 71)
combined part of Brachybelus with Pachyteuthis,
and the overall shape of the rostra (213) argte in
becauseofthe excentricity.The lattergenus(q. v.) has
favour of an associationwith the Passaloteuthinae. A
a clearly different juvenile rostrum, however; it
connection rnay then be sought via forms like
belongsto a much younger branch of the family. A
Psetrdohasrites(p. 234, Fig. 88e). (241) PerhapsB.
relationshipto Dactyloteulftlsis more likely.
t r a b e c u l aL i s s .( 1 9 1 5 ,P l . 1 , F i g . 7 - 8 ) ( L i a sy ) i s a n
B. breviformls (Voltz) is clearly a typical species.
intermediatespecies.
The following forms are more or less closelyrelated: B. zieteni (Werner) (:hreviforntis Zieten, cf. Fig. 85a),
The genusCylindroteuthriBayle 1878.
B. gingensis (Oppel), B. meta (Blainv., cf. Werner 1912, Pl. 12), B.
incuryatus (Zieten), B. brevis
Here belong speciesrelatedto B. puzosi d'Orb. from
( B l a i n v . ) ,B . i n s c u l p t u s( P h i l l . 1 8 6 4 ,P l . 4 a n d 5 ) , B .
the Upper Dogger, e.g. B. rediyiyus Blake" with
contrltrs(Rcimer),B. vulgaris (Young and B.), B. rttdis
c o n t i n u a t i o n su p t o t h e U p p e r N e o c o m i a n ( 8 .
(Phill. 1866,Pl. 16), B. "abbrevicttus"(d'Orb.,Pal. fr.
, s p e e t o n e n s i sP a v l o w ) . B . m a g n i J i c u sd ' O r b . , B .
j u r . , P l . 9 ) , B . " e r c e n t r i c u s " ( I b i d . P l . 1 1 ) , a n dB .
porrectLtsPhill. and B. obeliscusPhill. are important
crassusYoltz.
species.The rostrum is in generalslender-cylindrical and is characterizedby a rather shallow ventral groove
The genusHomaloteuthis Stolley 1919.
starting from the posterior end; it can become deeper due to corrosion.Unlike Stolley(ct p. 245), rve do not
B. spinattts Quenstedt(1858) and related forms
include here the Belentnopsis-like rostra with a deep
from the Lower Dogger belong here. They have a
ventral lurrow extending to the alveoh.rs (and
smooth, sharp,dorsally pointing apex without grooves
accompanyingits posterior part); theseare included in
or at most with slight traces of them. The juvenile
Aulacoteuthis.
rostrum seemsto be more elongate(Fig. 71) than in M e g o t e t r t h i s ; o t h e r w i s eH o m a l o t e u t f t i s c o u l d b e
The gennsPachyteuthisBayle 1878.
considereda subgenusof Megateuthls.We assumethat it was derived from Brachybelus, in the way that
B. excentralisYoung and Bird (1822)belongshere as
Megatetrthiswas derived from Odontobelus.
the type, along with its relatives (8. subcluadratus
(242)
panderianusd'Orb. and others),as well as the species
Rcinrer,B. lateralis Phill., B. explanatus Phill, B. d) The subfamily Cylindroteuthinae nov.
o f S t o l l e y ' s( 1 9 1 l ) A c r o r e u t h i s( F i g . 7 a ) . T h e s t o c k y
We consider Cylfudroteutil.r Bayle a typical genus of
shape of the adult rostrurr and its strongly excentric
this subfamilyand include in it especiallythe species
growth are similar to Brachybelu.s.The juvenile
which Stolley (1919) distributed between his three " f a m i l i e s " C y l i n d r o t e u t h i d a e ,O x y t e u t h i d a e a n d
rostrum,however,is clearly different from the latter, as
Pqhyteuthidae.In terms of their shapethey apparently
P a c ' h y t e u t h l s f i r s t a p p e a r si n t h e U p p e r D o g g e r ;
are relatively close to the Passaloteuthinae from which they differ by their juvenile rostra which are very
Quenstedt(1849) recognizedits peculiar features;he illustrated the excentric growth and the juvenile form
elongate.slender, club- or rod-shaped(Fig. 7ll-o). Suchjuvenile rostra also occur in the Belemnopsinae
$. 427, PI. 27, Fig. 5, Pl. 30, Ftg. 27) (cf. Keferstein 18 6 6 ,P l . 13 1 ,F i g s 15 a n d2 0 ) .
is the greater elongation of the adult rostrum.
and Duvaliinae, but these subfamilies are sharply distinguishedby special f-eatureswhich are lacking
The genusOxyteuthisStolley 1911.
here. The Cylindroteuthinae could either be descendants ofthe Passaloteuthinae. in which a rod- or
Accordingto StolleyB. brunsyiceilsls v. Stlorlbeckis
club-shaped rostrum was formed early and was
the type of this genus;it has a Cylindroteuthis-Iikc
subsequentlymodified, or they must be derived from
outline and no ventral grooves; it occurs in gt'eat
the hastites.A decision can onlv be expectedfronl
numbersin the Upper Neocomian.n'ithout any notable
t44
/1}
h , l:i't:'ll
v_/ ,.d
ilf '1
ffi
.,1
i] [.1]
$il tl a.
11Y ,..,H*, Fig.88.
Secondaryelongationand shorleningofthe rostrumin the generaSalpingotetrthis (a-d), Pseudohastites(e) and Cylindroteuthis (J-i).
a. B. acuaritts t,icanoliculatusafter euenstedt 184g,pl. 25, Fig. 13. Dorsal, lateral and ventral view. From the Lias ! ofHeiningen' aftcr Phillips. P1.13, Fig. 35, dorsalview; a2: the samein ventral view (probablyall ..rr:shorterspecimen;a2: B. qLtadricanaliculatus are identical). b. B. actrariuslongisulcatus(Quenstedt,Pl. 25, Fig. 33a). Lias t. br: crosssectionfor b (note inside:the old apicalfuirows). c. B. tripartitLts(Quenst..PI.26, Fig. 16a).Lateralview. c1:cross sectionof the apex. after Phillips (Pl. 12, Fig. 30). Ventral view. d1: dorsalview of apex. d. B. ilmensn.en.rl.s view of e. B. scabrosusafter Phillips (Pl. 20, Fig. 51) from the upper pafi ofthe Lower Lias (y?). Lateral view. e1:dorsal,ez:ventral apex. the juvenile stage.fi. (Same situationin ".8. owenii var. f. B. spiculaylsafter Phillips (Pl. 33, Fig. 82). Ventral view. Inside: Phill. Pl. 31, Fig. 76). puzosianus", pttzosi(d'Orb.) after Bayle, Pl. 25, Fig. 1. g. C1,lindroteuthis h. B. nitidus after Phi1l.,Pl. 13,Fig. 34, lateralview. /r1:crosssectionofapex, i2l crosSsectionofmiddle paft. the length of i. B. poryectusafter Phill. 1870,P1.32. Fig. 80. Anterior part somewhatshortened;i.e. must bc completedaccordingto the dotted arow. Samecomment as forf ,,Oxytettthis"or ,,Aulacotertlrls", posteriorhalf viewed from the right side (i.e. has to be completedanteriorlyby the samclength). k. After Stolley 1911,Pl. 8, Fig. 2. Note the courseof the doublelateralfurrows.
=
145 sign of speciation(Fig. 711, o). It differs from the
6tr.,Pl. 37,Pal. univ.,Pl. 77,Ft1.7). However,(246) |t.
s l e n d e rs p e c i e so f P a s s a l o t e u t h i s i n t h e j u v e n i l e
could also refer to the form shown in Figure 64c, the
rostrum.but the overall outline is similar. ln additionto
affiliation of
the typical species,Stolley(191l) citesa new O. pugio
Belemnoteuthis).
which is totally obscure. (cf.
and (245) O. jasikovi Lahusen(p. 178). The following
Rhaphibelus acicula is a strikingly small, thin,
sroupwas originallycreatedas a subgenus; in 1919(p.
markedly needle-shaped belemnitefrom the Solnhofen
5 I ) it was given the rank of a genus.
beds,without distinct furrows and with a circular cross section.Its overall shapeis reminiscentof the thin,
Tlre genusAulacoteuthisStolley ( I 91 1) 1919.
elongatespeciesof Salpingoteuthis(5. gracilis, Fig. 86b); it cannotof coursebe united with them since the
S t o l l e y ( 1 9 1 1 , p . 1 7 5 ) d e s i g n a t eB s . absolutiJbrmis
stratigraphicallevel is too ditferent. The absolutesize
Sinzow as the characteristicform of the genus; it
also counts.It suggeststhat we are looking at ajuvenile
diff-ersfrom C1;lindrotetrthis andOxvteltthistn having a
form that cannot of coursebelong to Salpingoteuthis.
deepand long ventral groove,thus the overall aspectis
But it could be the stage precedingthe club-shaped
similar to Belentnopsls.Under this name a genuswas
juvenile
createdby Lissajous(1915,p. 25-26);it overlapspartly
Belemnopsinaeor Cylindroteuthinaeof the Upper
with Stolley's genus. ln addition to the species
Jurassic.Fine specimensare housed in the Munich
mentioned,B. absolutusFischerv. Waldh. was placed
collections.
rostrum seen in
the
Duvaliinae,
there as the characteristicfonr.r.Stolley could probably plead againstsuch an extensionusing his arguments
(217)
formulatedin 1919 (p.52-56); but I do not think that
e) The subfamily Belemnopsinaenov.
he can rnaintain them all in their full extent. The
Here belong the "Basisulcati" Rcimer 1836, the
conformity of B. absolutiforntiswtth absoltttu.s must be
"Canaliculati"d'Orbigny 1842,Deslongchamps 1878,
noted, all the more so as there is a seriesof related
M a y e r 1 8 8 3 ,Q u e n s t e d1t 8 8 5 ,Z i t t e l 1 8 8 7 ,N e u m a y e r
speciesthat could easily be united here in the senseof
1890,Pavlou' 1892,or the "Hastatidae"Stolley 1919.
Lissajous.It is true that there are doubts about the
Moreoverthe "Belernnitellidae" Stolley1919.
position of several of them; they could belong to
The characteristicform of this particularly
B e l e n t n o p s i s , b u t t h i s c a n o n l y b e d e t e r m i n e di f
important and well known group is the genus
alveolar slits are shown to exist. To place them in
BelentnopsisBayle, the main characteristics being a
C,vlindroteutlris is not helpful in terms of clarity and
rod- or club-shaped,elongatejuvenile rostrum and a
systematicorderliness.Here I mention: B. gratfiianus
deep ventral alveolar groove in conjunctionwith an
d'Orb., B. sulcatus(Miller) Phill. Moreoverthe allies
alveolar slit. The shapeof the rostrum is very variable.
of B. blainvillei Yoltz (Fig. 89) need to be scrutinized,
Slenderconical forms occur along with rod- or club-
i.e. dpeciesthat Stolley regarded as belonging to
shapedones of different cross sectional outlines, the
Cylindrotettt i.r or to Belentnopsis(8. alpinus Ooster,
ventral groove can be limited to the alveolar end or
B.
extend more posteriorly,sometimesto the posterior
t r n i c a n a l i c t r l a t u s Z i e L e n ,B . s u b b l a i n v i l l e i
Deslongchamps, B. infracanaliculattrsQuenst.,as well
end. The alveoli (cf. d'Orbigny, cr6t., Pl. 5, Fig. 15)
as B.
n t t t n i e r i ,t e t r d m e r u s ) b r e y i c a n a l i s e t c . Deslongchamps). Nothing certaincan be said prior to
provide some information on the shapesof the pro-
investigationson the alveolar slit and the juvenile
to what we saw rn B. gigantee;s.ln addition to that we
rostrum.(cf. alsoSalpingoteuthis).
have some direct evidence:rostra of B. semisulcatus
ostracaand phragmocones;they give a picture similar
with remainsof the phragmoconeand impressionsof The genusRhaphibelusnov. gen.
the dorsal shieldsare preservedin Upper Jurassicstrata ( c f . p . 1 8 0 ) ; t h e y s h o w f e a t u r e so f A c a n t h o t e u t h i s
The belemnite from the Upper Malm shown in Figure
speciosa(Fig. 63), which may belonghere.Figure 90
9012belongshere.It may be identicalwith B. acicula
showsthe carefully reconstructedoverall picture of the
M t i n s t . 1 8 3 0( p . 8 , P l . 1 , F i g . 1 a ) ( a l s os e eK e f e r s t e i n
shellofa speciesofthis group.
1 8 3 4 ,p . 4 2 4 a n d d ' O r b i g n y 1 8 4 5 ,p . 5 6 7 , f 8 4 6 , P a l .
An essentialfeatureof the subfarnily is the ventral
146
v I
J j
utl
tl
,il {l i/I f
t
I"
! t .
I
Fig. li9. Moryhology of the Bclemnopsinae(r I nat. sizel ventralviews exceptg. dr. d2). a."Belemnopsis"unicanicultrtdHartm..afterBayle.P|.30. Fig.2, fiom Lcs Mouticrsncar Caen.with longitudinalfurrow dying out anteriorly(probablycloseiyrelatedto the following form). b. Belentnitesblainvillei after Phillips,PI. 25, Fig. 59, fiom Sherbomc. c. Shorter variety (.a-d probably belong to the cylindroteuthidslrAulac'oteuthist l, but they have to be comparedalso with SaIp i ngotetrIh i.s { Fig. 88 } andB elemnopsi s). d. B. strlcatusafter Phillips. Pl. 30, Fig. 75 (ct. p. 248). d2: crosssectionat the posteriorpart of the alveolus"r11:cross sectionin the areaof the incisionu'hich marks a distinct incuruingof the rostrallamellae. e. Belemnopsisbessina(d'Orb.) after Bayle, Pl. 30, Fig. 1. 9 "i g . 1 4 a . f l : c r o s s s e c t i o n o f t h e a l v c o l a r r e g i o n w i t h v e n t r a l s l i t ( o r i g i n a l ) . . [ : j u v e n i l e r o s t r u m J . B . h a s r c t mas f l e r Q u e n s t e d r , P l . 2 F after Quenstedt.Pl. 29. Fig. 35a. g. B. "subJirsiJbrntis" afterQuenstedt,Pl. 29. Fig. '13.Lateralview. h. B. bcudottinlaiter d'Orbigny (Pal. fr. t. crdt.,Pl. 5, Fig. 1),judging by the crosssectionit looks like a Belemnopsis.This form is so distinct that I shouldlike to createa special gents"Belemnoconns"ffor it], since anotherspecimenis known from the Neocomianof England(Dixon I 878. Pl. 27, Fig. 29). a-r./pcrhaps belongto lrlacoteuthis, the otherslo Belemnopsis(e) andHibolites (f, C').
"alveolar slit", a feature that can easily be studiedby
at the alveolar groove (218) are apparentlyinterrupted
splitting the shell medially (p. 200). In such a
a t t h e m i d - l i n e , o r a t m o s t a r e c o n n e c t e db y a n
preparationa typically limited field appearsin the split
extremelythin layer of shell rnaterial.The remainsof
surface,the so-called"slit field", which is very smooth,
the latter may be representedby the substanceon the
often coveredby a thin layer of chalky substancethat
"slit field", but that substancecould also have been
has sometimes been interpreted as an ostracal
taken up from the outside.Belemnitella mlrcronata
continuation of the conotheca extending into the
Schloth.shows this structurevery clearly (Fig. 70): on
rostrum. This interpretationis probably incorrect. (cf.
the inside of the intact alveolustwo lines run parallelto
D e s l o n g c h a m p s1 8 7 8 , w h o h a d a d o p t e d t h i s
the alveolar furrow; they could be regarded as
interpretationfrom Munier-Chalmas).An "ostracal
connectedwith the siphuncle,but in fact they belong to
larrella", as Neumayr (1889) called this supposed
the sheath(since the ostracumis smoothly detached),
structure,is not visible in Belemnop.slsand Hibolites.
and the wedge-like part lying in betweenthem can be
However,shell lamellae(Fig. 89f1)which are infolded
knocked off. The latter of course extends,as a shalp
147 Fig. 90.
Hibolites semisulc'atu.s (Milnst.). Reconstructionof the whole slrell. assumingthar Acanthoteuthisspeciosais identicalwith this species (as rs suggestedby contemporaneousoccurrenceand total agreement of stmctureof the phragmocones).phragmoconeshave rarely been found (in the lithographic limestonesof Bavaria and Su'abia) with the rostrum of Hibolites semisulcotus,somewhat morc olten with the pro-ostracum of Acanthoteuthis ,tpeciosa.The pro-ostracumis carefully drawn from the fine impression(slab and counterpartin thc Munich collections;this is the o r i g i n a l o f Z i t t e l . 1 8 8 5 ,p . 5 l l ) .
The phragmocone is drawn from an ercellent (three-dirrnensionally preserved)specimenin the Tilbingen collections,along with other specimensconfirming its associationwith the pro-ostracum.The rostrum n'as added on the basis of fine specimens(in the Munich collections)of Hiholites sentisulcatuswhich retain partsof the sheath(alveolus).The sipl.runcleis also addedon the basis of a specimen from the same collections. Thc only problematic point is the specific identity of thc phragmoconesin the two elements.which have never been found together in perfect preservation.Similar rostra and phragmocones are known fiom several specimens of Hib. hastatus. _ 1 : median longitudinal striae;2: a specialrydistinct growth rine of the median prate, which apparently demarcatesa delicate (uncalcified) marginal zone; J: boundary bctwecn middle and lateral plates at the growing margin; 4: asymptotic line, indicating the distance covered by the matrix during growth since a poinr conespondingto 3; 5: lateral plate with growth line; 6: anterior margin of annulus; z: antcrior suture line (the dark area is the last septum);,9:rostrum layer;9: ventral alveolarfunow; 10: crosssection in the posterior part of the alveolus; 11: young specimen with the impressionof the phragmocone(spccimcn liom Eichstiitt, in the Munich collections); l2: Rhophibetusacicula (Mtinst.) from Solnhofen (Munich collections),naturalsize.Needle-shaped rostrumand phragmocone.
edge or lamella, into the alveolar slit; it must be regarded as a distinct part of the alveolar sheath, correspondingto the so-called"ostracallamella',.An open slit cannot have been present here, and we thereforeare well advisedto nse neutralterms for these highly characteristicstructures.We shall call the iryerruption of the normal rostral mass in the median plane the "slit field", and its problematicfilline the "strafummedianum". It is noteworlhythat a complicationof this structure occursin a species(namelyB. sulcatusMiller from the Oxford Clay) which clearly belongs to the Belemnopsinae;this complicationpartly elucidatesand partly obscuresits character(Fig. 89d). The ventral groove deepens, slit-like behind the alveolus. Transversesectionsreveal (d1)that the growth lines are incurvedin the middle and are not closelyjoined to one another,so that longitudinal spacesresult below the groove.This situationseemsto suppoftmy inferenceof
148 penetratingligaments that causea perturbationof the
The genusHibolites Mayer-Eyrr-rar 1883.
tbrmation of the median rostral growth layers, thus controlling the whole differentiation of the rostrum in this zone. The shapeof the slit field and its correlationwith
Type: B. hastatusBlainv. 1827.Relatedspecies,some olwhich occur widely, are'.B. wiirttemberglcasOppel with weakly developeddorsai furrow; it seemsto be
the phragmoconeand rostrum exhibits wide variations.
the oldestHibolites (Bajocian,Dogger y). B. bel,richi
(cf. Fischer1887, In the oldestspeciesof Belemnop.sls
Oppel, B. helveticusMayer, B. latestrlcatttsYoltz, B.
p . 3 6 1 , F i g . 1 3 9 ) ,e . g .b e s s i n a t, h e s l i t f , r e l de x t e n d sa
semihastatusBlainv., B. planohaslalas Roemer, B.
long way backwards and slowly dies out towards the
girardoti
ventral groove. According to
Quenstedt, B. c'analictrlatusandB. hastatasshow the samecondition.
suhJusiJbrmlsRasp. The particular condition of the
In the specimens(249) of Hibolites hastatusthat I have
reduction, starting from the posterior end, already
examined,the slit field is limited to the anteriorpart of
begins in in the typical speciesof Hibolites (seeabove,
the rostrum.Its inner limit extendsposteriorlylrom the
F i g .7 0 ) .
L o r i o l , B . s e m i s t r l c a t u sM i i n s t . , B .
alveolarslit shouldbe studiedin individualspecies.Its
protoconchbut then disappears,and the slit field is
(250) As mentionedabove (p. 241), lt is possible
abruptly truncated(Fig. 70e) (cf. Quenstedt,Pl. 29, Fig. 29a). In the mesohibolitesand neohibolitesthis
thatHib. semisulcatus Miinst. (p. 180) andAc. speciosct
lirnit moves even farther forward (see these groups)
belemnite shell and animal in general, and ol this
and it reachesan extremeDositionrn Belemnitella and
family in particular,is greatly increased.We could then
Aulacoceras.
draw an almost completepicture of this specieson the
are identical.If this is so, onr detailedknowledgeof the
basisof direct observationeo. The genusBelemnopsisBayle 1878.
(251) The questionas to whether this identification is justified has been askedseveraltimes since Miinster
Here belong the Canaliculatrs. restr. Fischer 1887.
( p . 1 8 0 ) .T h u s H u x l e y ( 1 8 6 4 )w r i t e s :" A c a n t h o t e u t h i s
A t t l a c o b e l t t sP a v l o w 7 9 1 3 ,B e l e m n o p s i sL i s s a j o u s
speciosatlrrnsout to be one of the Belernnitidaebut the
1915.
statementsbefore us leave it doubtful, whether it was,
T y p e :. 8 . b e s s i n u ds ' O r b . ( F i s c h e r1 8 8 7 ,p . 3 6 1 ) t o
like Belentnoteuthis,devoid of an elongatedguard, or
which the known speciesB. canaliculallr.sSchloth.and
whetherit is really a Belemnite.s semisulcatuswith the
B.
guard broken off'.
apiciconus Blainv. are related. The oldest
This question was picked up by
representativeof the group could be B. harlel:i Mayer
Angermann ( 1902) without coming to a clear
from the Pliensbachian(Lias y), but this speciesis
conclusion.Let us quotefrom his text (p. 230):
problematic.Belemnopsis doubtless occurs in the lowermostMiddle Jurassic.
"Given the availablematerial of Acanthoteuthis, a n d o u r i n c o m p l e t ek n o w l e d g e o f B e l e m n i t e s
The rostra of these belemnitesare more or less
s e m i s u l c a t t t s , i t i s i m p o s s i b l et o p r o v i d e d i r e c t
slender-cylindrical,grooved along almost the whole
evidencefor the identity of these genera.We must
length (Fig. 89e). It is noteworthy that the juvenile
leave the questionfor future investigatorswho may be
rostra, at least of the later forms, are slightly club-
lucky in finding the evidence. What is certain,
shaped,in other words are Hibolites-like. (According
however, is that there is no reason to place
to Lissajous {1915, p. 23}, B. /usi/brmi.sParkinson shouldbe includedin B. bessina).As in the hibolites,
Acanthoteuthlswith the Belernnoteuthidae. According
the rostraare compresseddorso-ventrallyin the zone of
to our presentknowledgethe former could as well be a Belemnites semisulcatus."- That far Angermann's
the canal, and the alveolusalso appearstransversely
conviction based on similarity. And indeed, there are
oval in cross section. ln Belemnop.si.s the slit field
"important, if not compelling reasonsfor its truth, in
apparentlyalways extendsa long way backwardsand
that
gradually dies out towards the ventral groove
characteristicfeaturesof Belemnitessemisulcatus,one
(Quenstedt1849,Pl. 29, Fig. 5) (Fischer,loc. cit.. p.
of which is definitely absent,the other probably absent,
248).
in Belemnoteuthis":"The Munich collectionshousean
Acanthoteuthis clearly shows two very
r49 ,"1k!,,;
,a!, ir.il!il,
$lll."' d { 1d'ip' &,.rb
I 4,," l i f
il
u*d*F
#=ff -S * u**;x q9l 'l - S*+
%i,q ii; ,iir'i
dilp
,E*
:,.' rr *
"\
3
,*3,.'.i' ..,t .i$: i ,, ii rii
-"e,* Fig. 91. - An arm crou'n of "Acanlhoteutltisspeciosa" Miinst. Photographnatural size, after a slab from Eichstiitt (lithographic limestones.Upper Malm) in the collectionsof the Polytcchnic School at Braunschweig.This is the f,irstspecimenin which I have been able to establishthe presenceof 10 arms. One has to count the rows of hooks, some rows being without a correspondingarm impression.20 longitudinal ron's can thus be counted (see rows "1-20" marked on the figure). Most rows can bc traced from the numberedhook. and each ron can be distinguishedliom the others.Thc counterpartis in the Munich collections.On at least one specimenin the BavarianStateCollectionsI have been able to count 19 rows. - The impressionsof the headand mantle sac are vcry incornplete.Tracesof thc pro-ostracumare prcsent"and the ink sac is preserued.The arms are the importar-rt f-eatureof ttris invaluable slab;they are similar iu structurebut clitl-crcntin size.The two rows of hooks on eachaffn are alsounequallvdeveloped.
=
150 undoubted Belentnites semisttlcaluset from
the
beforeor during fossilization.For an originallyround,
Solnhofenbeds in which the impressionof the pro-
subsequently(during growth) dorsally grooved, and
ostracumis preserved" . "Acanthoteuthishas the sarne
finally again smoothly rounded structure of the
pro-ostracumas Belemnites semisulcatus" (p. 229).
phragmoconeseemsinconceivable.I think that these
Moreover the structureof the phragmoconeis virtually
pictures either representaccidentalchanges,or that
identicalin both fossils.especiallythe septalspacing
they are due to impressionsof the phragmocone,
which varies within the same limits. The sarnecan be
including the sheath,generatedduring fossilisation,
said of the apical angle. In the best preserved
after the dissolution of the periostracum(cf. p. 70
p h r a g m o c o n e so f A c d n t h o t e t t t h i s I f o u n d a n g l e s ranging from 20-22". The alveolus of a fine specirnen
ffootnotere],and Fig. 67a). This interpretationwould argue in favour of Belemnoteuthis,as would the shape
of Bel. semisulcatus had an angleof 20.3o.More acute
ofthe hooks(p. 186andFig. 68b,e).
anglesmay occur in both cases,but I only found them in poorly preservedspecimens.Even if identity did not
(2sl)
exist, the structureshown in Figure 90 would have to
The genusDicoelitesBcjhm 1906.
b e a s s u m e df o r H i b . s e m i s u l c a t u s O . nly the finer details of the pro-ostracum(253) would not represent
H e r e b e l o n g B e l e m n o p s i s - l i k ef o r m s c l o s e t o B e l .
directly observedfeatures.The soft body (p. 7) should
meyrati Ooster;in addition to a ventral alveolarfurrow
also correspondto the generaltype (Figs 62d, e,67a).
they show a dorsal one, apparentlywith a similar slit
The same applies to Ac. speciosa, which n-rust
f i e l d ( c f . B c j h n 1 9 0 6 ,p . 3 8 9 a n d L i s s a j o u s1 9 1 5 ,P l . 1 ,
additionallyhave had a typical belemnoidsheath.If the
Figs 2 and 8). The genus containsdoubtful forms (cf.
identity is real, then we have a belemnite specieswith
Stolley 1919,p. 44). Among the major representatives
rostrum, phragmocone,pro-ostracum,mantle sac, ink
Diener, D. waageniNeumayr.D. are'.D. .strlcacutus
sac, and head and arms including brachial armament
ketrwensis Bcihm, D. dicoelus Rothpletz. The group
( F i g s6 3 , 9 0 a n d9 1 ) .
appearsto occur widely throughoutthe Middle and
A minor doubt is raised by the existenceof a
U p p e r J u r a s s i c .H o w e v e r , t h e r e i s a p o s s i b i l i t y o f
peculiar groove which is faintly visible in the most
confusionwith normalhiboliteswith an inconspicuous
posterior part of the phragmoconein a few specimens
dorsal furrow (H. wtrerttembergicus Oppel?)or with
of Acanthoteuthis speciosa (in the Bavarian state
Duvaliinae(8. avena?).
collections, Munich), which is reminiscent of Belemnoteuthis antiqua(Fig. 67). But thesestructures
The genusMesohibolitesStolley 1919.
are not sufficiently similar and distinct to allow a different conclusion; perhaps they have been
Here are Neocomianspeciesclose to B. minaret Rasp.
accidentallyproducedin similar positions,as could be
( c f . S t o l l e y 1 9 1 9 ,p . 4 5 ) , w h i c h a r e a l s o k n o w n a s
expectedin this type of rock. In any case,this is a very
"Depressi".In addition to the speciesmentionedwe
questionable"distinctive feature" of the phragmocone
include
of Acanthoteuthis:indeed.while the groove in this parl
Schwetzoff and Hib. rhllgl Schw., perhaps also Hib.
of Belentnotetrthisbelongs to the periostracum,since
pingttis Schw., Hib. varians Schw., Hib. g,tgrictts
the phragmoconeor conotheca is round, we have a
Schw., Hib. /allauxi Uhlig and Hib. beskidenslsUhl. from the Upper Neocomian and the Aptian.
totally different situation lin Acanthoteuthisl:the periostracum is lacking (hence the uncertainty of
in
Mesohiholites:Hib. minaretiformis
Characteristicfeaturesare the marked shorteningof the
identification) and the slight groove is located on the
alveolar furrow and the slit field (p. 2a9) and the
conotheca, or rather on the steinkern of the
conspicuousdorso-ventral flattening, a very striking
phragmocone,sincethe conothecaitsell is not clearly
modificationof the hibolite type.
preserved.So in this respectthere can be no agreement with Belemnoteuthis. The question is whether new
The genusParahibolitesStolley 1915.
finds can show the typical occurrenceof such a groove in phragmocones of Acanthotettthi.s.lfso, one would
Stolley (1919,p.45) united in this genusa numberof
have to assume that it is due to a secondary effect
s m a l l s p e c i e s ( U p p e r N e o c o m i a n ,A l b i a n , L o w e r
l ) l
Fig. 92. - Schematicdrawings to illustrate the most common lorms of Aclinocamar afterCrick 190,1. u. A. verus Millerl b. A. granulatusBlainv.l c'. A. quadratus tl'Orb. (Gonioteuthis Bayle). The dotted parts of thc sheathor rostrum have been destroyedbefbre or during lbssilization (cf. ZitteI 1887, p. 507).
Cenornanian)and designatedP. duvaliaefomzl.rStolley
Closely related speciesoccur in the Senonian.with
as the type. They certainlyresemblethe Duvaliinaeand
more or less club-shaped,elongate (8. mucronatu,s
are often confusedwith them, given the strong lateral
Schloenbach1867,Fig. 2) or short cylindrical rostra
compression and deep, gutter-like double lateral
( 8 . h o e / b r i S c h l o e n b a c h1 8 6 7 "P l . 1 6 . F i g . 1 ) ; t h e y
grooves.Among them are P. pseudoduyaliaSinzow
differ from Neohibolites'in having very distinct.
1 9 1 3 ( : P . t o L r t i a e W e i g n e r 1 9 0 9 ) ,P . b l a n / b r d i
sometimesdeep lateral and vasculargrooves,and very
SpenglerandP. stoliczl'alSpengler(loc. cit. p. 46).
short alveolar furrorvs that end ah.r.rostsuddenly togetherwith the sharp slit. We agreewith d'Orbigny
The genusNeohibolitesStolley 1919. Here belongsa group of speciesfrom the Lower and M i d d l e C r e t a c e o u s( U p p e r N e o c o m i a n - U p p e r
( 1 8 4 5 )( s e ea l s oW a g n e r1 9 0 5a n d S t e i n r n a n ln9 l 0 ) i n emphasizingthe close relationsl.ripof this genus to speciesof Actinocanra.t.although."veleavethem rn a separategenus. Their branching vascular lurrows are
Cenomanian) with the type B. semicanaliculatus
less deeply incised, whereasthe lateral lines are of
B l a i n v . T h e y e x h i b i t a ( 2 5 5 ) s h a p er e m i n i s c e n to f
sirnilar form. The point added to the blunt, finger-
Belentnopsis,in that the club is only slightly, if at all
s h a p e d e n d a l s o c h a r a c t e r i z e st h e w h o l e g r o u p
swollen,whereasthe alveolarend appearsthick. The
("Mucronati"). A peculiarity of the alveoli in thesetr,,'o
ventral furrow is limited to the alveolarend and the slit
genera cannot be overlooked: they shorv an apical
doesnot extendbackwardsbeyondthe protoconch.It
angleof about20o,as in most Belemnopsinae, but are
has a similarlimit as in Belemnitella (Fig. 70, cf.
(Fig. 70). A roundedindentation slightly cor.npressed
Stolley 1919,p. 46). Concerningthis group seeStolley
on the mid-dorsalline showsthat a narrow,thickened
(1919: Die Hiboliten...)who figured a number of
median rib reinforced the rnedian plate ol the plo-
species(Pl. 1, Figs 1-32). Here we cite N. in.flexus-
ostracum (256) (or the conotheca).This structureis
gracilis Stoll. (Aptian), lV qff strombecki G. Mtiller
alsopresentin Actinocanrux.
(Albian), N. aptiensi.sKil., N. cf. ewaldi v. Strornb.,N. cf. ntinor Stoll., N. ninimus Blainv. (Fig. 83h). Some
The genusActinocamux Miller 1823.
Neohibolites shou'Actinocantar-like anterior ends which are due to corrosion.
A. verusMiller from the Lou,er Senonianand its allies fiom the Turonian and Middle Cenomanianbelong
The genusBelemnitellad'Orbigny 1845.
here. (Probably also Belentnocanto.rbov'eri Crick 1910,representative ofa groupbasedon small"perhaps
I n t h e p r e c e d i n gs e c t i o n su e h a l e r e c o g n i z e dt h e
juvenile individuals,which could be given the rank of
courseof evolutionleadingflom the Belemnopsinae to
subgenus).Theseforms are characterizedby a lessthan
Belemnitella, the type of rihich is B. tttucronata
solid constructionof the anteriorparts of the sheath,at
S c h l o t h(.c f . F i g . 7 0 a n d Z i r r e l! E 8 5 .p . - ; 0 8 .F i g . 6 9 8 ) .
least in the zone of the alveolus.which causedtheir
152
n
s
n nu,
K n
F ||
frreflu[i
\::t
glffif$ U ffit i,i p l ili, l lllll |il'tllrfit
Fig. 93.
t--/
size).
F:l
a. PsetrdobelLts bipartitus after d'Orbigny (Pal. fr. t. crdt..
l1:,:1 lrl; I
v f,,
Pl. 3. Fig. 6). a1:crosssectionofthc same. b. B. coquandtr.rd'Orb. (cf. Pal. Fr. jur., P1.21). br, br:
l l li '
d,n0
correspondingcrosssectionsfrom a specimenin the Ziirich
,,.,l.j
(Polytechnic School) collections. no. ./ 2626 from the
&:i I rff i
Argovian near Chdtel St. Denis. Labels read: Ha,stite,s
gil
l$,1
ltil
0u,
cll V
fn", U
o vl
LJ'r
c. B. exili.s fiom the Lias e fl-ower Toarcian] after
l
A
trf
t,
f"
\t!2
N
souvanatri d' Orb.,Pseudobehrs monsaIvensis.
e.R_l .x
t{ J
\lt I
m b ,t V
v
Rostra of Duvaliinae and similar types (r/2 nat.
i.\\r/
1
Pl. 25, Fig. 16. Latcraland ventralviews.c1.c2: Quenstedt. correspondingcrosssections(cf. p. 228).
,n
tl. B. polygonalls Blv. after Bayle. d1. d2: corresponding cross sections, from diff-erent spccimens. Upper row d: lateral view of the same speciesafter Quenstedt 1849. Pl.
n,Wr
30, Fig. 9. da: ventral view of the same.betweend and d, the dorsal view, and cross sectionsof the alveolar and
rostralpafts. Specimensfrorn the Neocomianof Castellane(cf. p. 258). e. Conoteuthisc'onophorafrotr the Tithonian ofthe Strambergbeds,after Zittel 1868 and 1883 combined.Anterior part in ventral, postcriorpart in dorsalview. with dorsalfurrow and tangentialview of thc alveolus. sameafter Bayle, Pl. 31, Fig.l3. /. Dtrvalialata at\erPictetand Campiche1858.Pl. 13, Fig. 10a.y':conespondingcrosssection..l3: /a: sameafter d'Orbigny, Pal. fi. t. crdt.P1.4,Fig.5. f1: D. dilatota (ibid. Pl. 3. Fig. 3). g. Ibid. aflerBayle"Pl. 31. Fig. 14. h. Duvalio entericiRasp.After the figure in Bayle,Pl. 33. lateralvicw. l7: anteriorview of fracturesurface1r11nat. size),.r:dorsal kee1.
post mortem destructionand normal loss. Insteadof an
( 1 8 8 7 ) a n d P a v l o w ( 1 8 9 2 ) a t t e m p ts u c h a g r o u p i n g
alveolus we find a pseudoalveolus,which is much
under the name "Dilatati", without arriving at a sharp
wider and of variable cross section,but the destroyed
definition of a natural group. Clearly related forrns
part sometimesextendedfarther back, so that a conical
were excluded(e.g. Bipartiti, Conophori).The rnost
or pyramidal anterior end of the rostrum resulted
striking featureof the Duvaliinae is the dorsal alveolar
insteadof a pseudoalveolus, the concavepart of which
furrow, whereasa ventral furrow is missing. There are
once contair.red the protoconch.The easily-destroyed
several additional features of the habitus with
parts are well defined so that the resulting forms of
somewhat doubtful weight, which neverthelessmake
corrosionare characteristicofthe differentspecies.
up an unusual picture: the deep lateral furrows with
It was once assumedthat Actinocamax and certain
sharp double lines, the lateral compression,the strong
species of Hibolites or Neohibolites,which are
dorso-ventralasymmetryof many species.The juvenile
normally cor-rodedat the anteriorend (without showing
rostra are elongate-cylindricalto club-shaped,at least
the same regularity, cf. p.
in the speciesof Duvalia.It would be interestingto see
202), never had
phragmocones. This of courseis out of the question.
the correspondingfeaturesof the oldesttypes relatedto
D i s t i n c t p h r a g m o c o n er e m a i n s a r e k n o w n i n l .
Pleurobeltrs (Fig. 851) and Rhabdobelus (Fig. 93c).
quadratus. See Figure 92 for the structureof normal
(258) Derivation from theselorms is indeed suggested
forms.
by the shape.
(2s7)
The genusDuvalia Bayle 1878.
I) The subfamily Duvaliinae (Pavlow) em. HerebelongbelemnitesrelatedIo Duvalia Bayle,some
Bayle (1878,Pl. 21, Fig. 3) distinguishes the "Dilatati"
o f w h i c h w e r e d i s t i n g u i s h e da s " N o t o c o e l i " b y
of other authors(Deslongchamps,Zittel) as a separate
d ' O r b i g n y ( 1 8 4 2 ) . D e s l o n g c h a m p st 18 7 5 . ) , Z i t t e l
genus. The type is Bel. latus Blainv.; the most
r53 distinctive featureof the subfamily is the strong lateral
extinctorius(Rasp.).
cor.npression. For other charactersthe speciesincluded, e . g . D . d i l a r o t o ( B l v . ) ( s e eZ i t t e l 1 8 8 5 ,p . 5 0 7 , F i g .
g) The subfamilyBayanoteuthinae nov.
695) are highly variable;they may be elongateor short,
Here belong the genusBavanoteuthlsMun.-Chalmas
almost knobbly, sometimespointed, sornetimesblunt
and its allies from the Eocene, which are perhaps
ended.Figure 93f shows a typical profile. An extreme
related to Belemnitella, but external features
form with a dorsal keel (x) is shown in Fig. 93h
distinguishthem from the latter (internal charactersare
(Dtrvaliaemericii Rasp.).It is difllcult to imagine a
not known in detail). The phragmoconesare extremely
biological role for such bizarre structures in a
slender.
belemnite rostrllln. (The shortening could be interpretedas an indication of a return to a littoral life
The genusBayanoteuthr'.s Mun.-Chalmas1872.
style). I considerthe different, very peculiar forms of B. pol.vgonallsBlainv. (Fig. 93d) to be somehow
The genus includesB. rttgi/br Schloenbach1867, a
related to Dut,alia, but they should probably be
speciesthat is strikingly reminiscentof aulacoceratids.
accommodatedin a distinct genus,Pseudoduvalian.
The very slender, cylindrical rostrum is pointed
gen.
posteriorly, like Cvlindroteuthis. But it shows fine longitudinal grooveswhich cover the whole surface,in
The genusPseudobelusBlainville 1827.
a more irregular pattern than is shown in our Figure 94f. In the area of the alveolus there are broad dorso-
T h e t r u e " B i p a r t i t i " ( D u v a l - J o u v e1 8 4 1 ,Z i t t e l 1 8 8 1 , P a v l o w , 1 8 9 2 ) - P s e u d o b e l u sP a v l o w l 9 l 3 b e l o n g
the pear-shapedcross section grades into a sub-
here.Type: Pseudobelusbipartitus Blainv. 1827 (p.
tetragonalone. The alveolus is strikingly slender(9o);
lateral furrows, which soon die out posteriorly, so that
113).In additionto the slightdorso-ventral asymmetry,
in this zone there is no ventral furrow nor a slit.
there is a distinct feature in the considerabledepth of
Apparently related to Bayanoteuthisis St-vracoteuthis
the double lateral flrrow.
(260) Crick 1905, with a stocky, cylindrical-conical,
which results in the
"bipartite" aspect of the rostrum. This is already recognizablein Rhabdobelusexili.sd'Orb. (Fig. 93c)
biunt sheathwith a deep alveolus (12') and ventrolateral alveolar furrows (a mid-ventral one 1s
but is much more distinct in medium to large sized
uncefiain).
specimens of bipartittts.Somemeso-and parahiboiites (8. coquandusd'Orb., Oxfbrdian,Fig. 81b, cf. d'Orb.,
h) Review.
Pal. fr. jur.. Pl. 21) show the samephenomenon almost
I am not yet in a position to follow the evolutionary
equallyclearly.
series,speciesby species,throughthe Mesozoicstrata. I must be content to hint at the general relationships
The genusConobelusStolley1919.
betweenthe subtarrilies.After this prelirrinary work, a comprehensivetreatment of the known material by a
Lissajous(1915) createdthe genusRhopaloteuthis for
careful, stratigraphicallyand morphologically trained
some slender. only slightly (lateraily) compressed
researchercould hopefully "read from the rocks" an
"Conophori" (Mayer-Eymat,Zittel 1883).At present
important part of history. The visual grasp of external
this genus(259) rs hardly distinguishable.According to
form should be supported by careful structural and
Lissajous.B. satn,anatrid'Orb.shouldbe the type, and
developrnental studieswhich could be achievedusing
related forms are B. gilieroni Mayer, B. spissus
split specimens and polishedmediansections(Fig. 71).
Gilidron andB. conophonr.s Oppe| (cf. Zittel 1885,p.
Since any belemnite rostrum allows one to read its
506,Fig. 694).Hou,ever"the flrst two speciesprobably
ontogeny,we have extremely useful material here for
belongto the Hibolites(cf'.Ba1'le1878.P|.29, Figs 5-
seriousmorphogeneticstudies,which we were unable
7. and d'Olbigny,Pal. fr. jur. Pl. 2l). B. conophL,rtrs, in
to utilise fully for want of time and rleans. At least
contrast, is a well defined" Dttyalitt-llke form that
some baselinesof synthetic description have been
(Stoll*' 1919.p. 49). establishes the genusConobeltr.s
drawn, however, indicating the future direction of
H e r e t o o b e l o n g C . o r b i g n r n t t t r . r( O p p e l ) a n d C .
research. A very general question is that of the
154 Fig. 94. - Belemnoidsfrom the Eocene('i2 nat. size). a. Vasseuria occ'identalis reconstructcd. F
ffi1t L$ c.
a
&. The same.large specimenafter Cossmann1895-98,Fig. -:-t:! e
10. c. Alveolus opened longitudinally,strikingly blunt (cf. p. 2 81 ) .
,"F\ /'=?-\
i::" 1
\{+/
t ! t
d. Striationinside,ventral. e. Striation inside, dorsal. Parabolic lines and hypcrbolar zone(longitudinalstriation).
\_.-/ @ r(s)) /
relationship between the oldest known belernnites
J. BelemnitesrugosLtsafter Schloenbach1868, with distinct dorso-lateralfurrow. 1-4: respectivc positions of the cross sectionsin g. l. Longitudinalsectionwith the very slenderalveolus.
Thesebelemnoidshells(Fig. 65a-c),which occurrn
(Nannobelus andHastires), which is especially
the black shales(Wengenbeds near Raibl, Oberloch)
important (p. 225) becauseit affects hypothesesabout
of the alpine Triassic,show very striking peculiarities
the derivation of the younger "clavirostrid" types. The
which are still problematic. A rostrum proper is
latter certainly are not as uniform in characteras
apparentlylacking, but the situationcould be similar to
assumedby Abel (1916), but vague similarities of
that in Acanthoteuthisspeciosa(p. 251); the sheathis
certain forms to Hastitinae,which were mentioned
not preserved,sometimesthere are at best sometraces.
above more than once. are undeniable.In addition to
The phragmocone is rather blunt, the sutures are
the club-shapedjuvenile rostrum they appear with a
inclined forwardson the dorsal side.Really noteworlhy
more slenderrostrum, rnore widely spacedsepta,and
is the pro-ostracumwhich appearstripartite,in that the
longer septalnecks; in all these aspectsthey appear
lateral plates are drawn forwards on either side of the
closer to
typical median plate; there is only a narrow separation
the
alllacoceratids than
do
the
Passaloteuthinae.
in the fom of zoneswith concavegrowth and marginal lines. Of the soft parts the ink sac is preserved; moreoverthere are impressionsof the cephalicorgans,
E. The family PhragmoteuthidaeNaef l92l
which can hardly be recognized in detail, and the
(System,p. 534).
typical double rows of small hooks indicatingthe arms. Suess(1865) placesthe speciesin Belemnites,but he
The very peculiar structure of the pro-ostracum
believes (without any good reason)that the anterior
requires the creation of a special group for the
part of the phragmoconecontains mere "ligations"
follou,ing form:
(supportingledges)but no septa.In contrast,he had a clear idea of the pro-ostracum.Its greatpeculiarityis
(261)
only recognizedwhen one attemptsto reconstructit. I
The genusPhragmoteutftlsMojs. 1882.
first despaired of integrating such a shell into a decapodbody; then I thought of a possibility of trying
Here belongs a single known species:Phr. bisinuata
(In it on the basis of the recent genusThysonotettthis.
(Bronn).
the latter the shell has laterallobes,but they are curved
S y n o n y m sB : e l e m n o t e u t h ibsi s i n u a t aB r o n n 1 8 5 9
into the body where they support the visceral mass
( J a h r b . )p,. 4 4 , P I . 1 , F i g . 1 - 3 ;A c a n t h o t e u t h b i si s i n u a t a
laterally, adjoining the funnel retractors.Cephalopoda,
s i s i n u a t aK e f e r s t . S r i s s1 8 6 5 ,P l . 1 - 4 ;A c a n t h o t e u t h i b
vol. 1, chapter37). However, Figure 67 now shows
1 8 6 6 ,P l . 1 3 , F i g . 5 - 7 ; A c a n t h o t e t r t h i sb i s i n u a t a
how a normal integration with the shell of
Mojsisovics1902 (Cdphal.,p. 199 where small hooks
Phragmotetrthis can be imagined. One has to assume
and the pro-ostracumof a special species {?} are
that the "dorsallobe", i.e. the medianplate of the pro-
describedand frgured)(cf. Fig. 68d).
ostracum,did not extend (262) to the very end of the
=
155 mantle,and that the whole pro-ostracum,especiallythe
latter. But even though the generalappearancemay
lateral plates, was delicate and flexible, as already
suggestthis possibility,a closer look at the essential
suggestedby the fossil specimens.Their elasticitymay
lacts shows that the aulacoceratidscorrespondto the
then have helped in opening up the mantle cavity
belemnitesin all the essentialfeatures;they undeniably
during the respiratoryand locomotory pulsations,as
have the characterof dibranchiates.Thereforeoossiblv remainingdoubtswill be eliminatedbelow.
can be assumedto happen with the broad vanes of recentLoliginidae(Figs 7c and 58b). Still, the genushas a very isolatedposition within the Belemnoidea;this is noteworthygiven the fact that it representsa very ancient type, which existed along with the aulacoceratidsbut showeda different extreme
The phragmoconeis indeed comparableto that of a r n o d e r a t e l ys l e n d e r O r t h o r : e r a s : t h i s s i m i l a r i t y i s emphasizedby the (compared to belemnites) r'ery widely (264) spaced septa which are srrong and
(rather than an undifferentiatedprototype). Shells
markedly I'aulted, and by the thick conotheca.The lengthof chambersin generalisr,2 to r,1of their width.
resemblingthe hypotheticalprotodecapod(Fig. 62)
The siphuncleis thin, moderatelynarrou,edat the septa
occur in Europe only in the Lower Lias, in the form of
(Fig. 95f), inflatedbehind them. It lies ventrallyas in
the belemnitesNannobe|usandCoeloteuthi,s.
a l l o t h e r d i b r a n c h i a t e s .( H a r - r e r1 8 6 0 a n d m a n y
Shells resemblingPhragmoteuthis have not so far
subsequentauthors clairned a dorsal siphuncle for
been found in other strata.Huxley (186a) describesa
A u l a c o c e r a s ) .A t t h e i r o r i g i n s t h e s e p t a ln e c k s a r e
phragmoconefrom the Lias with very conspicuous,
drawn slightly anteriorly, but they are not as a whole
apparently similar growth lines (Fig. 65d); closer
directedanteriorlye2. The phragmoconesare alrvays
inspectionreveals a belemnite cone, however. This
very slendercomparedto other belernnoids.The apical
suggeststhat the diverging. feather-likelines on either side of the middle stripe have to be interpreted
angies vary lrom 5 to l2'. I lbund apical angles ranging l0-11' in "Belentnites"spec.(housedin the
according to Figure 63b, i.e. as an imprint of the
Munich collections)frorn Malm strataat Streitbergand
./batheredpattern of the pro-ostracum(cf. Zittel 1883, p . 5 0 1 , F i g . 6 8 4 ) . T h e p h r a g m o c o n ed e s c r i b e db y
from Acanthlcas layers at Brentoniconear Rovereto (Atractites?).
Huxley could well belong to a normal Passaloteuthis
As in Spirula and the belemnites.the protoconchis
or C.vlindrotettthis,as envisagedalready by Huxley
roughly globular and
himself. However, this cannot be safely claimed (8.
appearsinflated comparedto the secondchamber.The
b r u g h i e r i ? ) . H u x l e y h a d n o p r e c i s ei d e a y e t o f t h e
conothecaclearly showsthe thicker nacreousand the
typical outline of the pro-ostracum.
thinner polcelain layer; both are smooth, devoid of longitudinalridges - in contrastto what is generally
F.
The
family
Aulacoceratidae Bernard
18950s. restr.
given its greaterdiameter
s a i d !( c f . v . B t i l o w 1 9 1 5 ,P l . 6 2 { 6 \ , F i g s 3 a n d 4 . yA .t best the upper most layer of the ostracum shows delicatelongitudinallines. As far as I have been able to seein specimensfrom coliectionsand also in Btilon"s
To
our
knowledge,
the
aulacoceratids
(Aulacoceratides)have been treatedas a family fbr the first tirre by Bernard(1895,p. 683) who also included
fine figures, the first delicate longitudinal ribs lwhere they occur) already belong to the periostracum,as do the subsequentenforcing layers (Fig. 95i); that they
the genusXiphoteuthis. They are olten considered a
could be derived from those of the orthocerans
subfamily of the Belemnitidae;a closer analysisof the
thereforeis out of the question(cf. p. 266').The rib-1ess
features of both groups does not yield the sharp distinction hoped for by systematists,since there are
atractitesare older! The posteriorpartsof sheathsof the aulacoceratids
s i m i l a r i t i e sp o i n t i n g i n b o t h d i r e c t i o n s ,a s n i c e l y
are reinforced.as in the belemnites,and they extend
demonstrated by Steinmann(1910): in additionto the
beyond the phragrnoconeas short tips ol as cylindrical
similarity to belemnitesthere is an evident (263)
to club-shapedprocesses.Only theseparts should be
relation to the orthocerans;thus one might assumean
called rostra (s. str.). But in order to clarity the commonly(265) vagueutilisationof the term (p. 175),
intermediary group between the tetrabranchiatesand the dibranchiates,ratherthan a definite affiliation to the
we have enlarged the concept and here again
t56 Fig, 95. , *^wt,r.r 1,/
l ii(\
t\
-4. .^
,
n
'
The sheathand rostrum in aulacoceratids(1/2nat.
s1ze.).
\e rt-
1 l,'
a. Dic'tvoc'onites reticulatu.r. Sheath with thc enclosed phragmoconcin dorsai view. Note the fine, regular fluting
li /S
./S o .-"J"
and the curvedgrowth lines on the anteriorpart (p. 265). D. Rostrum of the samespcciesin left lateral view. Note the longitudinal rib above the lateral fumow. Next to the main figures:crosssections.A1l after Mojsisowics 1902(Pl. 14). c. Rostrum of Atractites haueri with alveolus,aftcr Branco 1 8 8 0( P l .2 0 ,F i g . 1 ,p . 4 0 1 ) . d. Rostrum and alveolusof Aulacoceras ,sulcatum,spit horizontally (along lhe latcral furrows). After v. Biilou,' 1915,Pl. 57, Fig. 3. One can recognizethe normal growth lines, a distinct axial thread, and along the lattcr. in the posterior part, a radially fibrous structure as in belemnites (Fig. 7 I ); moreovcr one observesa sort of flbrillar structure radiating fron.rthc posterior end of the alveolus (cf. p. 278, and v. Biilow. Pl. 58, Fig. 3). A typical featureis the slender phragmocone.the posteriorend of u'hich is shown in natural sizenext to the main figure. e. Median section of a different specimenafter Btilow, Pl. 5 8 ,F i g . 6 . ./. The same continued and (exactly in the rnedian plane) completedschematically. g. Whole specimenviewed from the left side, after Biilow, P l . 5 7 .F i g . l a . /r. Crosssectionof anteriorend of g (Biilow, Pl. 57, Fig. 1c). l. Cross scction closer to posterior end, from a different specimen(Biilow,p. 38, Fig. lUb).
ft. Crosssectionof the anteriorzone from a similar specimenof Dic\oconites cf. haLteri(Biilow, Pl. 59, Fig. 9d1. /. Crosssectionof Dlcr. plonus (Bi.ilow,Pl. 60, Fig. 4d).
distinguish"rostra" having a large or a small alveolar
sheath, but rather on its inside in the form of
part. The latter (Fig. 95) is inconspicuousin the
i m p r e s s i o n s( F i g . 6 3 d ) . W h a t W a n n e r ( 1 9 1 1 ) a n d
actually club-shapeddictyoconitesand atractitesand
before him Mojsisowrcz (1902) interpretedas growth
the club thus can representa rostrumproper.
lines in fossil shells (Fig. 95a) is clearly a different
The growth of the aulacoceratidsheathsdoes not
sculpture,which may have been formed during the
really differ from that of belemnites.Only the layering
forward movement of the shell epithelium over the
is not as easily observed as in belemnites; in the
growing shell; it is not of specialinterest.The growth
rostrufir too the regular alternation of dense shell
lines of the conothecamust have been coveredrapidly
iamellae and radial substanceis not easily observable.
from behind; they indeed have a different aspectthan
When looking at these structuresit is important to
t h e s ec u r v e dl i n e s( c f . B t i l o w 1 9 1 5 ,F i g . 1 2 - 1 3 ,p . 2 6 -
remember:setting aside the protoconch,each point on
21). h is rather inconceivablethat the free rrargin of
the conotheca lying inside the rostrum was once
the shell had the outline of thesemarkings.They swing
situated at the anterior end and carried only a thin
forwards both ventrally and dorsally (though more
sheaths. str. All rostral structuress. 1. (p. 175) are
weakly here),laterally they are concaveforwards;there
secondarily superimposed on
those primary
is no hyperbolar zone, and a tongue-shapedpro-
differentiations.Moreover: The growth lines of the
ostracumof considerablelength, as is likely in such
conotheca,representingthe free shell margin of earlier
slender belemnoids, seems incompatible with such
stages,should not be sought on the outside of the
growth lines. The growth lines proper of the conotheca
r57 I r r re n o t 1 e tb e e no b s e r v e idn t h i s l a l n i l y . ln the aulacoceratidsthe rostra show distinct lateral
predecessors and closerelativesof the belemnites(cf. especiallyAtractites).
lirrlows, as in the belemnites,and there is no reasonto intelpret them differently (p. 198). Unfortunately they
Q67)
have been termed "asymptotic furrows", in belemnites,
The genusAulacocerasv. Hauer 1860.
too. (Apparently the markings on the sheathhave been confused with (266) the underlying markings on the
Theseare the best-knownaulacoceratids surrounding
conotheca).They have different features;as a general
t h e t y p e A u l . s u l c a t t r n lv o n H a u e r 1 8 6 0 ,a s w e l l a s
lule they are displacedventrally in the posteriorpart
Asterocon itesTeller I 885.
fof the rostrum], whereas anteriorly they lie more dorsally.beneathsu'ellingson the sheath.
D i a g n o s i s ( a c c o r d i n gt o B i i l o w 1 9 1 5 . p . l 8 ) : rostrum elongate,straight,club-shaped.drawn out into
To have an idea of the aulacoceratidanimals,we
a terminal spine,with very strong,straightlongitudinal
c a n o n l y a t t e m p t a r e c o n s t r u c t i o nf o l l o w i n g t h e
ribs which begin above the terminal spine and extend
nrethodsexplained fbr the belemnites(p. 186). We
to the upper end. From the apex a strong longitudinal
have to imagine these animals as very slender
furrow extendson either side to the upper end of the
dibranchiates.The Aulacoceras presentedin Figure
rostrum. There the furrow is less distinct than at the
95g would have rneasuredabout I m, in the size l c d u c t i o n u s e d f o r t h e i l l u s t r a t i o na b o u t I 2 r n : t h e
lower end. The phragmoconeis long and has a small apical angle (5-12'). Septa very widely spaced.
dictyoconitefigured next to it would have measured
Siphunclemarginal, ventrale3,fully calcified in its
about60 cm.
upperparts.
The aulacoceratids beganin the Lower Triassicat
This diagnosis is misleading due to a lack of
the latest(in the Penr-rian accordingto Steinmann1910;
zoological and morphological background; the
cf. Haniel 1915) and continuedat least to the Upper Triassic(Atractites).Their older remainswere mostly
following critical remarks are necessary:the "lower"
degraded during fossilisation,hence our rather
proper (cf. p. 175), i.e. the post-alveolarpart of the
incompleteknowledge.
sheath,is short and barely swollen (Fig. 95d), it is in
end is the posteriorend of the animal. The rostrum
The earlierhypothesesput forward on the ontogeny
fact pointed. What are available as fossils are the
and phylogenyol the aulacoceratids lack a sufficient
posterior ends of phragmoconeswith their thick
morphological,especiallyontogenetic, basis.This also
sheaths,not merely rostra, with inconspicuousalveoli,
a p p l i e s t o B r " i l o w ' s( 1 9 1 5 , p , 3 a 4 7 ) d e s c r i p t i o n .
as is the generalrule in belemnites.The "upper end of
Neverthelessthis work reflects al1 that can be said
the rostrum" is thus the anterior portion of the
positively.Negativepointsare: the longitudinalribs of
preservedpart of the phragmoconesheath;we do not
the aulacoceratids have nothing to do with those of
know how far anteriorly it originally extended. I
certain orthoceratids;they are at most analogousto t h e m ( p . 2 6 4 ) . T h e a u l a c o c e r a t i d sw e r e t r u e
neverthelessconsiderour Figure 95 as instructive and
dibranchiatesand had an internal shell from the earliest
parts were situatedbehind a slenderconical part, in
ontogenetic stages; there is no reason to assume
which the sheath became progressivelythinner
assumethat the cylindrical and ciub-shapedterminal
anything different. The assurnptionof dorsal and
anteriorly,similar to what is known in belemnites.The
ventral mantle lobes is refutedon p. 13. In Sepia the
latter part must have been destroyed prior to
shell doesnot arise as a "simple calcareouspart" lying
fossilisation.
at the surfaceof the mantle (cf. Figs 38 and 60). In
Growth proceeds by concentric increments,
terrnsof its form. Vasseurlais not closely relatedto the
revealedby grinding (Fig. 95d) as a fine layering (268)
a u l a c o c e r a t i d s ,e v e n l e s s s o i f w e i n c l u d e t h e
reminiscentof that found in belemnites.In contrastto
(c1.p. 60). mysteriousBelo.sepiella
the latter, however,we see no distinct alternationof compact lamellae and fibrous substance.- Biilow
We imagine the ontogenyof an aulacoceratidto be like that in other dibranchiates(Fig. l0c) and regard
(1915) observeda structurewhich is definitely not
the piesent family as a variant of the decapodtype of
comparable to the radial fibrous structure of the belemniterostrum. This structure(Fie. 95d) radiates
F i g u r e 6 2 . l n p a r t i c u l a r w e c o n s i d e rt h e m t o b e
-
158 outwards,forwards and backwardsfrom a zone which
pafts are met. The lolds of the shell epithelium had to
lies closeto the posteriorend of the alveolus;Biilow's
line increasinglynarrow grooves until they had to
(p. 35) interpretationis probably correct:the shell fold
withdraw frorr the filled ones. The secretionof shell
(B. calls it the "mantle") grew anteriorly and posteriorly along with the rostrum, thus enlarging its
substanceon the rudimentary ribs of the sheath (cf. B i i l o w , F i g . 1 5 a n d P I . 5 9 , F i g . 1 , s h o w i n gt h e -
z o n e o f i n c r e m e n t a li n c r e a s ei n b o t h d i r e c t i o n s .
erroneouslyinterpreted irnpressions)forms prismatic
"During this growth in opposite directions one part necessarilyformed a 'dead point', and this seemsto be
and fibrous structures lying perpendicular to the
the position at which the two directions of growth
is foided, less than perfect median sections(Fig. 95d)
balanced each other". Starting from this point the
do not show uniform structures, in contrast to
feathered radiation was formed due to continuing
transverse sections(Btilow's Fig. 15).Thereareno ribs
growth. This means that we cannot talk about
on the terminal spine; therefore the radial librous
homology with the radial structure of belemnites,
structure appears here in its simple fonn, as in
which otherwisehave strictly comparablestructures:
belemnites.(As regardsfiner surface sculpture see v.
In Aulacocera.s(Fig. 95d) we alreadyfind fibrous
surface,as in the axial part (p. 268). Since the surface
Btilow,p. 25, Figs 9, 10).
structurein the region of the apical line (which is like
The lateral furrows of Aulacoceras are strikingly
that in the belemnites{p. 204} and suggestsa similar
unequal: the dorsal one is not always distinct from
morphological interpretation);this fibrous structureis
other intercostaliurrows; the ventral one forms a long
especially distinct towards the posterior end and
depressionincorporating severalribs. Both are absent
demarcatesa roughly conical, axial structurelying between the terminal spine and the protoconch.
from the terrninal spine;they begin togetheras a broad depressionwith coalescinglines on either side, just
C o n s i d e r i n g t h e w h o l e d e v e l o p m e n t a ls e q u e n c e
anterior to the spine; anteriorly they gradually die out.
readablefrom the structureof the rostrum (p. 265), this
Between the two iurrows lies a longitudinal ridge
apical part is simply the portion correspondingto the
which is particularly swollen in the middle part of the
terminal spine at earliergrowth stages;there is no point
rostrum, forming a projecting roof above the ventral
in comparing this structure with an "embryonic
furrow. The dorsal lateral furrow appears to have
rostrum" or a juvenile phase of the rostrum, as Abel
a c c o m m o d a t e dl a r g e v e s s e l s ; i n d e e d g u t t e r - 1 i k e
( 1 9 1 6 p, . 1 3 1 - 1 3 3s)u g g e s t eadn d S t o l l e y( 1 9 1 9 p, . 1 3 )
imprints of vesselsleave this furrow, especiallyin the
subsequentlyaccepted.To understandthe details one
middle and in the posterior parts of the rostrum. They
shouldnote:
often show dichotomous branching and are probably
The growth of the rostrum in Attlacocelas
is
complicatedby the surfacesculpture,for the successive
homologousto those rn Dictyoconlle.r,which are more distinct(p.210).
layers have to foilow the fine and coarseornamentof
Growth lines of the conothecaare unknown (p.
the surface. Apparently the first additions were very
265). On the other hand it shows delicate longitudinal
fine longitudinalribs separatedby broad guttersor thin
ribs lying in the direction of growth, as in belemnites.
intermediatelayers;they can be seenat the anteriorend
More conspicuousis a broad ridge which is visible in
of preservedsheaths(Fig. 95h) even in adult stages
the mid-dorsal(270) line of a steinkern;it has been
(although they then appear coarser), and also in the
called "normal line" (as in nautiloids and ammonoids)
(269) deepestiayer (Fig. 95i) in more posterior
(Biilow, p. 32, Fig. 16). It proves that there was a
positions
whether in polished sections or rn
shallow groove on the midline of the conotheca,
preparationswhich have removed the outer layers
probably correspondingto the median rib of the pro-
(Btilow 1915,Pl. 57 and 58). Ribs of this form may
ostracum(cf. figurep.201: d1).
have been presenton the (lost) anterior part of the
The material studied by Biilow, to whom we owe
phragmocone.In the posterior part the ribs inevitably
most of our knowledge,belongs to Atrl. timorense
become thicker and broader, so that the grooves between them turn into sharp furrows. This
Wanner, a speciesclose to Aul. sulcatum v. Hauer, which - accordingto Biilow (p. 17) is a mere variety.
developrnentcan be followed by following the anterior
Other species rvhich probably belong here are
part of the sheathbackwardsso that increasinglyolder
A s t e r o c o n i t e s s a v u t i c u sB c i h u ra n d A u l a c o c e r a s
159 .lturlottenseWhiteaves(1889,p. 149),as well as some
we imagine them as suggestedp. 266. It is indeed
.)t' the typeslisted from Sicily by Gemmelaro( I 904).
striking that the vascular imprints on the rostrum
The genusAulacocerasis restrictedto the Triassic r Ladinian,Carnian,Norian).
resemblethose observedin the youngestbelemnites (Steinmann1910).But one should not pay too much atter.rtionto such features.They are less notervorthy
Tlre genusDictyoconitesMojs. 1902.
than the fact that Mont Blanc. when looked at from a certain direction, appearsto show the profile of the
Tlre aulacoceratidsrelated to Dict. reticulatus Hauer
great Napoleon.Indeed, vesselsrun along the rostrum
rc'semblethe genusAulacoceros in having at leasta
and split into branches a fact that is due to the role
I'e',vstrong, lateral, iongitudinal ribs on the sheath.In
this part plays in the organisationof all belemnoid
contrastto the latter they have an elongate,club-shaped
animals. Why they left strong or weak or no
post-alveolarrostrum. Due to the lateral extensionsit
impressionscannotbe explainedtoday; in fact. it is not
appearsdorso-ventrallyflattened(Figs 95a,b, k, 1).We
e s s e n t i a lt o k n o w i t . I n a n y e v e n t , t h e r e r s n o
also know, from some specimens,the anterior part of
signilicant "typical similarity" or "form relation"
the phragmoconeand its sheath.The forrner resembles
betweenDictvoconitesandBelemnitella.
thatof Aulacoceras(angle:5-10'), the lattershowsfine longitudinal striation, which partly grades into a
The genusAtractitesv. Ci.imbel1861.
legular fluting. In the group of the "striatl" (Mojs.), i.e. the subgenusDictyoconitess. str. Mojs.), this fluting
Rostraof aulacoceratids devoid of longitudinairibs and
also covers the posterior part of the rostrum. together
vascularfurrows are placed in the genusAtractites,
rvith vascrilarirnpressions reminiscentof Belemnitella
along r.l'ith large phragmoconesfrom the same strata,
(cf. Steinmann1910,p. 109).In the "laeves"Mojs.. i.e.
someof wl.richarevery largeindeed(0.5-1m). In some
( 1 9 1 0 ,p . 1 1 5 ) t h e s u b g e n uA s c t i n o c o n i t e .Sst e i n m a n n
instances^ the latter show distinct paraboliclines on the
this sculptureis missing. Here only the lateral furrows
dorsal side (Zittel 1885,p.456). flanked by strongly,
and the accompanying swollen ridges adorn the
longitudinally striatedfields; these structurescould be
rostrum. The lateral furrows differ from those of
the only (uncertain)evidencefor a pro-ostracumin the
Atrlacoceras:the dorsal one is deeply impressedinto
aulacoceratids. We note that the lateral plates of
the surface and in the posterior region (rostrr.rmand
belemnitesalways show a longitudinal striation due to
alveolarend) is directedtowardsthe midline; anteriorly it increasinglyacquiresa transverseorientatior.r. The
straight,delicate rlbs (272), so a sirnilar interpretation is justified here. - In any casethese structuresdeserve
ventral one is a shallow, often broad gutter. which is
further detailedstudy; after all the situation is perhaps
either smooth (271) or shows line longitudinal striations.The lateral ridge separatingthese furrows is
the sameas in Aulacoceras(p.265), the paraboliclines belongingto the sheath,i.e. to its innermostlayers.-
variable in thickness.lt can be a strongly projecting
These phragmocones are either circular or oval
ledge (Fig. 95a, 1) or may becomeinconspicuousand
(laterally cornpressed) in transversesection.lnside they
ahnostinvisible(k). Lateralridgesalso occur dorsally
show the typical structureof aulacoceratids;but there
and ventrally to the zone of lateral fumows,which may
are forms in which the septa are sorneu,'hat more
be united into a single furrow; these lateral ridges
closely spaced(chamberlength smallerthan chamber
extend from the sheath to the rostrum, so that three
width), thus coming closerto the belemnites,to which
strongribs on either side characterizethe overall aspect
in all eventsthey must be related.
of dictyoconites.
Of greater significance are the details of the
For individual speciesand finer structuressee v.
structure of the rostra. First one has to note their
Biilow (1915). I must again criticize his tendencyto
diversityin outline:the crosssectionrnay be circular,
(loc. cit. derive theseforms from certainorthoceratids
rarely somewhat compresseddorso-ventrally.More
p. 44).Likewisethey cannotbe cornbinedwith certain
often it is an upright oval, the narro$'estpart olwhich
belemnitesT . h e s h e l l s o f d i c t v o c o n i t e sa r e w i d e l y distributedin the Middle and Uppe-rTriassic(Camian-
is directeddorsally;it can also becomesub-quadratic with a slight depressionon the lower side. - The
Norian levels).As to the featureso1-thc lir ing animal.
rostrum in profile may be a moderatelyslendercone
160 with a sizeablealveolus,or a pointed cylinder, or even
Salzkammergut,Austria) is an interestingintermediate
a club that is in general compressed laterally,
form. But we do not wish to relate it closely to
representingthe rostrumproperwith or without a small
Pachyteuthiserplanata (Bull.) of the Upper Malm; we
alveolus. Including the preserved parts of the
s e e i t a s c l o s e rt o N a n n o b e l u s a n d B r a c h y b e l u s ,t o
phragmoconesheathsuch rostra may measure0.2-1 m
which the whole form (profile, cross section,
in length, attaining the thicknessof a human arm (cf.
excentricity)can easilybe compared.Nevertheles,we
Steinmann1910,p. 117),so that a total lengthof 4-6 m
do not wish to prematurelyobliteratethe limit between
can be assumedfor the whole animal. Such forms of
the aulacoceratidsand the belemnitids,becausethis
atractitescannot be consideredthe ancestorsof the
would be detrimentalto an objective assessment of
earliestbelemnites.In contrast,there are also forms
t h e i r t y p i c a l r e l a t i o n s h i p .I t r e r n a i n sq u e s t i o n a b l e
which haverostrathe size of a little finger.
whether the hastites(cf . p. 225) can be directly related
On the sidesofthe rostra one often finds dorsaland ventral furrows; but they are frequently indistinct and in some specieseven invisible. When they are fully
to the club-shapedatractitessuch as A. tenuirostris Hauer. The genusZugmontitesReis 1907doesnot belong
differentiatedone can recognizethe following features
here.It has short,markedlyblunt phragrrocones,which
(Bnlow 1915, p. 54): the lateral furrows are widely
are slightly curved in the siphunculardirection,with
spacedand differ in aspect;the dorsal one is rather
closely spaced septa (from the Triassic: Anisian)
broad and shallow, becoming deeper only on the
(Phragmoteuthtdae? Phragmoceras?).
anterior part of the rostrum, whereason the posterior part it curves slightly ventrally in the middle of the
The genusCalliconitesGemmellaro1904.
club, grows broaderand finally dies out. The ventral furrows are only delicate lines, and the distances
Unfortunately I
between them are srnaller than the distance from the
. e ep . 3 1 0 ,P l . 2 4 , F t g . 1 6 , G e m m e l l a r o n l y r e c e n t l yS
dorsal one. They are straight and extend farther
Pl. 30, Ftgs 23-26 in his work. Given the indicationsof
posteriorlythan the dorsalone.
an apparentreductionof the phragmoconeby Diener in
(273) Whereas the surface of many atractites is
received the publication by
the FossiliumCatalogus(1915,p. 23), v. Btilow (ibid.
totally smooth, others show a delicate granulation,
1921,p. 75), and (274)Broili (Zittel,Grundziige1921),
r e c o g n i z e db y B i i l o w ( 1 9 1 5 ,p . 5 5 ) t o b e d u e t o t h e
I had assumedthat this was a transitionalform leading
internal structure:when the rostra are well preserved,
to the teuthoids(Cephalopoda, vol. I, p. 135;cf. above
they show the radial fibrous structure of belemnite
p.104).
rostra,although it is sometimesindistinct and never as
Calliconitesdieneri Gemmellaro 1904 is a true
conspicuousas in the belemnitesproper. The "fibres"
a u l a c o c e r a t i do f t h e U p p e r T r i a s s i c o f S i c i l y ,
are formed by the finest crystals of aragonite, and
apparentlyrelated to Atractites. The rostrum has a
where they reach the surface they causethe granular
smoothor finely granulatedsurface,is slender,pointed,
elevations.
cylindrical to club-shaped,and laterally compressed.It
Atractites occur from the lowermost Triassic
shows widely spaced dorsal and ventral, iateral
(perhapsPermian, p. 264, 266') to the Upper Lias.
furrows. The phragmoconeis located in a slightly
Given the diversity of their form and their morphologicalsimilarity, they - especialiythe short, small ones among them - can be regarded as
excentric alveolus and shows the typical aulacoceratid Fig. 16) shows that the septa are extremely crowded,
predecessorsof the Belemnitidae.In particular we
and accordingto PI.30, Fig. 17 a long living chamber
might look here for the common ancestorsof the
occupies most of the rod-shaped cone following
N a n n o b e l u s a n dH a s t i l e s t y p e s . B u t c u r r e n t l y
numerous,very nalTow chambers.
form (loc. cit. Pl. 30, Fig. 25). Another figure (Pl. 24,
inadequateknowledgedoes not allow us to consider
(The relationshipof the isolatedphragmoconeswith
the specieslevel. The indicationsgiven by Steinmann,
the rostra seemsvery questionable,however; I think
based on overall similarities, will therefore not be criticized here. We gladly admit that Atr. quadratoides
they are orthoceratids:the apical angle is only about 3o, 1/r, the chamber length only its width. cf. loc. cit. Pl.
Steinmann (Upper Triassic "Rcitelstein" from the
3 0 ,F i g s 1 6 - 2 2 ) .
161 A problematiccase.
The phragmocone is built like that of an
H e r e w e m a y ( c a u t i o u s l y )m e n t i o n L a n g e r h a h n ' s
auiacoceratid,very slender and strikingly small in
( 1 9 0 6 , p . 4 2 ) " B e l e n t n o t e u t l , i l s s" p e c . ,f r o m t h e
proporlion to the other parts.The rostrum also seemsto
Triassic.lt is a spoon-shaped structurewith a peculiar
be rather rudimentary,it has a blunt posterior end,
shagreensurface texture, which is interpretedas a
without clearradial hbrous structure.
dorso-ventrally flattened rostrum (our Fig. 65k). The
Relationshipto Atractites is problematic: I do not
anterior part seems to grade into a sheath with an
excludethe possibility that Huxley's pro-ostracumis in
alveolus that shows traces of chambers.Given the
fact the rostrum of an aulacoceratid.An illusion of this
unusual characterof the "rostrum". it is uncertain
k i n d i s c o n c e i v a b l e .I n H u x l e y ' s s p e c i m e n s o f
whether this interpretationis correct. The available
Xiphoteuthisthe club-shapedanteriorthickening is not
indicationsare definitelynot convincing.If it could be
in onepiecewith the rest of the fossil.
studiedin greaterdetail,the natureof the phragmocone (?) misht be clarified.
Branco (1880, Zeitschr.f. g. N., p. 401, Pl. 20)
On the other hand, I note the following facts: l. describesAulacocerasliassicum(Giimbel sp.) from the lowermost Lias of Liimmerbach near Salzburg and
G. The family XiphoteuthidaeNaef 1921
gives an illustration (Figs 7 and 8) of a compressed
(System,p. 53a).
specimen.This specimenis housed in the Bavarian State Collections under the name Atractites liassicLnu.
The following genus and specieshas such an unusual
Along with it I saw specimensfrom Kammerkahrthat
character that it
shorveda very peculiar aspect.(276) | am under the
cannot be united with the
aulacoceratids.
impressionthat one of them resembledthe spindleshapedanteriorend of a largeXiphotetllrzsHuxley, and
(27s)
I indeed supposeit is the same species.Mojsisowics
The genusXrphoteuthisHuxley 1864.
( 1 9 0 2 , S u p p l . ,p . 1 9 8 , P l . 1 6 , F i g . 2 ) a l s o f i g u r e sa fossil which he interpreted as a rostrum of Atractites
Including "Orthocera elongata" de la Beche : B. tl1act'oconus Kurr, - Orthoceratites macroconltsKurr,
spec.;Steinmann(1910,p. 117) illustratesit againand
: O. liasinu,sFraas.- B. orthoceratoidesFnren.
rostrum has the same character.(Compare it with
calls it Atractites applanatusSteinm.This (Norian)
I can confirm that the speciesoccurs in the Lower
Atractites ausseeonltsMojs. and considerMojsisowics
Lias of Englandand Lorraine(specimenfrom Malroy
1902,p. 199,Pl. 15,Fig. 2). Could the pro-ostracum of
near Metz, provided by Krantz {Bonn}, now in the M u n i c h p u b l i c c o l l e c t i o n ) .l t s m o s t c h a r a c t e r i s t i c
Xiphoteuthis be merely an erroneousaddition? Or do
featureis the very long pro-ostracumwhich is spindle-
question cannot be answered yet for lack of
shaped,narrow posteriorly and thickened anteriorly
comparativematerial. Since in any event Xiphotetrthis
(Fig. 66). If Huxley's observationsare correct (see
(given the structllreof the phragmocone)is closeto the
below) and the situationis really as he describedit, we
atractites,it must be placed with them (cf. Keferstein
have to accept a very remarkable structure,which
1 8 6 6P , l . 1 3 l , F i g s l 0 - 1 2 a n dZ i t t e l 1 8 8 5 p , . 4 9 6 ,F i g .
played the role of a backbonein a special way in this
617).
the atractitesindeed possesssuch pro-ostraca?The
very slenderanimal. If so it cannothave been made of heavy shell material. which would have been concentratedin the ver1,area where gas is located in other types (Sepia,Ascoceras).It seemsmore likely
H. The family Belemnoteuthidae(Zittel 1885)
that the thick pro-ostracurnwas a light structure,
(System,p. 534).
Naefl92l
p e r h a p s e v e n c o n t a i n i n g - e a s .O n l y t h u s c a n w e understandHuxley's specimen.namely as the shell of a
The 1-amilyis basedon the charactersof the sheathin
slendernektonic-pelagicbelemnoid.The crystallized
the following genus and species,which is doubtless
substancenow filling the plo-ostracum spindle is
closeto the belemnites.
doubtlessa productof fossilisation.
= =
t62 The genusBelemnoteathisPearce1842.
C o e l o t e u t h i s ( p . 2 3 1 , F i g . 8 4 i ) . T h e d i s t i n c tr a d i a l
Including B. antiqua (Cunnington): Belemnotetrthis
structure of the rostrum and sheath (cf. Quenstedt 1 8 4 7 , P l . 2 6 ) p l a c e s B e l e m n o t e u l / z l sc l o s e t o t h e
antiqua
(Cunn.) Pearce 1842:
"Onychotettthis
Belemnitidae, from the oldest forms of which it may
antiqua" Owen 1844,p. 65-66 (also "Belemnosepia", "Acanthoteuthis"let- Belemnitespuzosiantrs d'Orb.
possiblybe derived.As tn Coeloteuthis the rostrum
1 8 4 5 ,P l . 3 4 , a n d 1 8 4 6 ,P a l . u n i v . , P l . 3 5 a n d 5 6 : Belemnoteuthisantiqtra Mantell 1848 : Belemnites
The very restricted load on the phragmocone suggeststhaLBelemnoteuthiswas a surfaceswimmer.
o v , e n i i Q u e n s t e d1 t 8 4 9p . 4 3 6 , 5 2 5 ,P l . 3 6 ( F i g . 1 9
In making the reconstructionI have deviatedfrom the
therein actually showsa belemnitewith ventral furrow) : Acanthoteuthis antiqua Morris 1885
originals on certainpoints: I have addedfins oftypical
Belemnoteuthisantiqua Woodward 1856 (cf. Zittel
decayedmaterial lying besidethe mantle as fins. I have
1 8 8 5 ,p . 5 1 2 , F i s c h e r1 8 8 7 ,p . 3 6 5 , F i g . 1 4 3 , a n d
also completed the arms and the eyes using typical
Quensted1 t 8 8 5 ,p . 5 1 0 , P l . 3 9 { " O n l , c h o t e u t h i s owenii"\,\.
details (edge of the lid, swimming membranes),thus
The
literature contains much misleading
showsno concentriclayering.
form (Fig. 62), rather than interpreting the displaced
producinga conceivable,life-like dibranchiate. This type is limited to the Oxfordian fCallovian]; it
information on the highly interesting species
has been found in the "Ornatenton" of Christian
Belemnoteuthisantiqua (Cunn.) Pearce,due parlly to
Malford
confusion, partly to premature interpretation of the
(Wiirttemberg). The fine phragmoconesfrom the
numerouswell-preservedfossils. (277) They show the
secondlocality have an apical angle of 2l-22'. (cf. p.
phragmocone,the sheath with a short rostrum, the
251 on Acanthoteuthisspeciosa).Septalspacingis also
muscularmantle, ink sac, head with eyes, funnel, and
roughly the same as in the posterior part of the
arms with hooks. Apparently no traces of the pro-
phragmoconesof B. semisulcatusandAc. spectosa.
ostracumhave been found (?). But seeHuxley (1864)
The possibility of a close relationshipto the nearly
who describesit, perhapson the basis of erroneous
lc. speciosais thereforestill worthy contemporaneous
According to Fischer(1887, Fig. l43a) observations.
of consideration.We have here a definitely belemnite-
the middle of the dorsal side shows distinct parabolic
sheath. like animalwith an underdeveloned
(Wiltshire) (278) and Garrmelshausen
lines giving at least an indication. We have no doubts that a pro-ostracumof the kind shown in Figure 90 is present;the delicatenature of the structuremay hinder
I. The family Diploconidae nov.
its preservationand thus preclude its appearance. (Unfortunately I have not yet seen the fine original
The genusDiploconus Zittel 1868.
specimensfrom England). We have here a belemnoid species in which direct observationspermit a full
The following species is now separatedfrom the
r e c o n s t r u c t i o na, s i n t h e c a s e o f A c a n t h o t e u t h i s
belemnites,from which it probably originated (as did
speciosa (Bel. semisulcatus?p. 250). The resulting
Belemnoteutils), becauseof its strikingly narrow pro-
conceptionsof the general type are mostly conflrmed
ostracum (Fig. 65i) and the absence(?) of radial
(Fig. 67b); a new investigationof the original
fibrous structure in the rostrum, which moreover is
specimenswould probablyrevealmore details.
s h o r t a n d b l u n t : i t s t r u e p o s i t i o ni s n o t y e t c l e a r .
The outline of the sheathis consideredtypical; but
however.Diploconus belemnitoidesZirtel 1868 cannot
its texture is strikingly fragile and the transverse
be combined with any other known genus; it was
section shows some special characters: a fine
merely to avoid further monotypic families that I
reinforcing rib lies on the ventral midline, flanked by
(1921) left.Diploconers in the Belemnoteuthidae(see
inconspicuousfunows; on the dorsal side there are two
the new edition of Zittel's Grundziige, edited by
swollen ridges, on either side of a median groove,
Broili). Perhaps we have here a transitional form
which diverge anteriorly before gradually dying out
leading from the belemnoidsto the sepioids(cf. p. 3l),
(Fig. 67). The rostrum is a shorl, slightly roundedapex
as suggestedby the short, bulky sheath, the strong
of a cone. the outline of which is reminiscent of
ventral curvature and its earliest occurrencein the
r63 uppermostMalm (Tithonian).For detailsseeFig. 65f-i, which was drawn from the original specimensand Zittel's figures. A remarkablefearureis the slanting septa, their upper side being curved forwards at the mid-dorsalline. This is partly causedby the oval cross section of the phragmocone,the oval outline being drawn out like the pointedend of a hen's egg.
Diploconus ZitLel,if one disregardsthe strong rib (cf. Fig. 70d) at the mid-dorsal line, i.e. a c o n s p i c u o u s thickening of the narrow pro-ostracum. In any event one is looking at a belemnoid type that might be includedhere. (cf. Kefersrein1866,Pl. 1 3 0 ,F i g s 1 4 1 6 ,F i s c h e1r 8 8 7P , l .2 , F t g . 9 ) . (?) The genusAmblybelus n. gen.
The genusConoteuthisd'Orb. 1842. Here belongs C. dupini(anu.s) d'Orbigny
1842 (Annales),Pl. 12,Figs 1-5,p. 377 (PaI.fr. crdt.,pl. t ). c f . a l s o 1 8 4 5 ,1 8 5 5 p , . 4 4 4 , P l . 3 2 , 1 8 4 6( P a l .u n i v . ) ,p t . 30. Owen (1844) placed ConoteuthiswjthBeletnnite,s. as did Zittel (1885) and others.Quenstedtregardedit as an "On.vchoteuthis".As the sheathis tnknown (279.
A problematic lbssil should be briefly recalledhere; it has (280) already been mentioned(p. 205). namely B. o b t u s u sB l a i n v . ( F i g . 7 1 v ) . I f t h e i l l u s t r a t i o n sa r e a t ieast partly correct (which is rather doubtful), they might indicate a Coeloteullrl.s-likeanimal. They show fairly pointed phragmocones(20-2l.) with closely
only small phragmoconesfrorn the Aptian of France a n d ( a c c o r d i n gt o W o o d w a r d1 8 5 6 ,p . 4 0 2 , 6 0 1 ) f r o m
spacedsepta.In contrastto all other belemnoidsthe protoconchappearsbroad and totally flat (as in certain nautiloids),making this specimenvery problematic.
the Gault ol England, the systematic position is problematic. Ail that we know about this peculiar
The sheath has a belemnoid structurebut does not show a distinct rostrum; it merely forms a slightly
belemnoid is the (not very satisfactory)information from the author. The specimenis unique; apparentlyit
thickenedenvelopearoundthe peculiar initial part of
was damagedduring study and now seemsto be lost. D ' O r b i g n y ( A n n a l e s , p . 3 6 6 ) s t a t e s :" A m o n g t h e
specimendescribedby Knorr was housedin "Klein's
numerous!rmpoftantcommunicationsthat I owe to Dr. Dupin, there was a small cone to which this hardworking naturalistdrew my attention.He had found it in the Upper Neocomianclay near Ervy (Aube). At
the phragmocone. Thereis no apicalline. The original collections"; it is said to have been found near Lauffenberg.(It is uncertainwhether Blainville had seenthe specimenhimself).If the figures are correcr, they clearly representa separatebelemnoid genus,fbr which we suggestthe nameAmblr-belus.
first sight I took this cone for something like the alveolarcone of Belemnites.At close examinationI first recognized a more arched form, with more obliquely orientedsepta;and with a good lens I saw the growth lines of the horny shell imprinted on the pyritic alveolus, indicating - not a broadenedshell as in Belentnites - but a very narrow one, analogousin all aspects to the ommastrephids. I call this form
K. The family VasseuriidaeNaef l92l (Systemp. 63a). As with the previousexamples,this family owes its isolatedposition to the impossibility of placing the following genuscloseto any otherform.
C o n o t e u t h i s ; l i k e a b e l e m n i t ew i t h a n a r r o w s h e l l similar to that of Ommastrephes,or an Omntastrephes with an alveolarconelike that of Belemnites".
The genusVusseuriaMun.-Chalmas1880.
D'Orbigny's figure (loc. cit., Pl. 12, Fig. t) of this supposed intermediate form is of coLrrse a
This is one of the few belemnoidsfrom the Eocene, and appearsto be widely distributed in France (paris
r e c o n s t r u c t i o nc, l o s e l y ( a n d w i t h o u t g o o d r e a s o n ) approaching an ommatostrephid.Indeed we know
basin, Loire, Brittany): Vasseuria occidentalis Mun.-
nothing about the relative length of the pro-ostracum, likewise the free margin of thc conushas never been s e e n .D ' O r b i g n y ( 1 8 4 5 ) s a y s h i r n s e l f t h a t t h e n o n chamberedpart has been added to the figure. What remains are the features of a phracmoconeof
C h . 1 8 8 0 .p . 2 9 1 . S e e t h e f i g u r e sa n d d e s c r i p t i o nb y V a s s e u (r 1 8 8 1 ,P l . 1 , F i g s 8 - 1 5 ) ,a n d F i s c h e r( 1 8 S 7 p ,. 359, Fig. 137), as well as the description of Belosepiellagiven earlier(p. 60). The nraterial cornprisessheathsof Vasseuria, of a Dentalium-like shape;in superficialaspectthey are
=
r64 L. Review of the fossil Belemnoidea and their eYolution. The Aulacoceratidae are probably the oldest belemnoids,occurring from the Lower Triassic, or
&# Fig. 96.
"Rostrum and phragmocone of Ilasseuria
even from the Permian(p. 266).I have not been able to extract sufficient information on their stratigraphic distribution from the literature, on which to base geneticconsiderations. They were probably derived from the orthoceratids,but they already representtrue dibranchiates.Their relation to the belemnites is
occidentalisMunier-Chalmas".From Abel 1916,p. 146, Fig. 60. - In fact only the upper part belongs to Vasseuria,
through the genus Atractites, which ranges to the
whereasthe lower part is a Belosepielloaddedarbitrarily (cf.
Upper Lias, thus permitting a connectionin time. A
p. 60).
more detailedsequenceof transitionalspeciescannot be given at present.Perhapsthe belemnitesoriginated from aulacoceratids via two stem groups.This is at least suggested(282) by the early separation(p. 225
certainly reminiscent of Aulacoceras.Thereforethe presentgenus has recently (again) been regardedas a p. 37, descendant of this ancientgenus(v. Biilow 1,915, Steinmann 1910, p. 107). The coarse,rather (281) irregular ribs and shalp grooves of the sheath indeed suggestsuch a relationship.But that is about all that remains after a closer inspection:the alveolus (Fig. 94c) is very short and blunt, and somewhat shifted ventrally as in belemnites.Judgingfrom the remainsol suturesthe septamust have been oblique in orientation, which might suggesta relationshipwith belopterids.In additionto the suturesthe inside of the alveolusreveals lines that can be interpretedas growth lines. They curve forward dorsally in an arcuatefashion (parabolic iines), and this zone is limited laterally by longitudinal areasthat may hide the hyperbolar zones.The alveoli are too small and erodedto reveal such fine details.On the ventral side one can make out arcuatelines which are concave forwards, so that an interpretationas the free shell margin is likely to be cor-rect(Fig. 94c, d, e). In Fischer's figures the hyperbolar zone is not (loc. cit. p. 359). represented Our conceptionof the bearer of these sheathsis given in Figure 94a. We see there a belemnoidanimal characterizedby a very slendershapeand a relatively heavy rostrum which grades directly into the thick sheath;the rostrum is slightly curved dorsally. Given the somewhat irregular form of the sheaths, we consider Vasseuriato have orobablv been a bottomlivins belemnoid.
and 258) of the Hastites and Nannobelustypes,which are not clearly linked to one anotheramong the earliest belemnites.In the Upper Lias the earliestmembersof the other subfamilies are already linked to the P a s s a l o t e u t h i n a ei,. e . t h e p r o t o t y p e s o f L i a s s i c belemnitesclosely related to Nannobelers.(At least Belemnopsis and Cylindroteuthis). Here again a detailed evolutionary seriescannot yet be worked out. The origin of the generaStyracotettthis and Bayanoteuthlsfrom the Eoceneis quite obscure.The latter is reminiscentof Cylindrotetrthis,whereasthe former may be more similar to Belemnitel/a.The small belemnoid groups can be regarded as very minor brancheso.f the main stem'.Phragmoteuthis from the Triassic is completelyisolatedfrom the aulacoceratids, Xiphoteuthis of the Lower Lias is in a problematic position in relation to Atractites.Belemnoteutlls of the Middle Jurassic could be a belated descendantof Coelotetrthls.LikewiseDiploconus could be a descendantof ancient belemnites of the Nannobelus type; its structureand its occurrencein the uppermost Upper Jurassicsuggestthe possibility of a relationship with the sepioids, which appear in the Eocene. Vasseuriacannotbe closelyplacedanywhere.
165 (283) P a r t V : T h e O c t o p o d a oor octopus-like
extremities of the transversallyextended,unpaired shell rudiment which lies within the muscularmantle.in which the inner face of each arm bearstwo rows of
dibranchiates. (p. 28a). Contents: A. Diagnoses.B. Generalconsiderations
cirri lying on either side of, and alternating with the suckers,which are arrangedin a singlerow, - in which
C. The genusPalaeoctopus (.p. 2 8 5 ) . D . T h e g e n u s
the arms are connectedthroughoutmost of their length
.lrgctnauta(p. 287).
by a very largevelar membrane.Typically planktonic. 2. Polypodoideaare octopodsdevoid of trr.refins (which may be replacedby lateral skin ridges or folds), - in which the arms bear 1 or 2 rows of suckerswhich are never accornpaniedby cirri, - in which the velar
A. Diagnoses.
Octopods are dibranchiateshaving eight arms of generally similar form; the third pair of arms in
membranegenerallyleavesthe greaterpart of the arms free, - in which the shell rudiment is representedby
octopods,numberedfrom above,is hornologousto the
two separate,cartilaginousrodlets, or is completely
tentaculararms of decapods,the dorsal pair being absent,- in which the cups of the arm suckers,which
lacking.Typically benthrc.
are arrangedin one or two rows, are not reinforcedby a horny ring so that the suckerscannot be transformed into hooks (!),
B. Generalconsiderations.
in which the suckers are not
demarcatedby a deep constriction frorn the muscular
The octopods are discussedhere only for the sake of
"carrier" or basal cushion, although they rnay appear "stalked"when the carrier is strongly extended,- in
completeness.Since their shells are, and probably
which a buccal arm crown or "buccal funnel" is
very scarceremainsfiom past geologicalperiodsat our
entirely lacking,
disposale5.
in which the funnel tube always
lacks a funnel valve,
always were, extremely rudimentary,we have only
in which the renal pores are still
We do know (Voltz, 1835, Appelldf, 1899) that
associated on eithersidewith the gill base,namelythe
octopodsform an internal shell anlage;they always
proximal part of the efl-erentbranchial vessel,- in
producean early shell sac rudiment that is similar to
u'hich the dorsal mantle margin is broadly fused with the head,a nuchal plate being absent,- in which the
the correspondingrudiment in decapods,but which is smaller than that 1n (285) decapods("Cephalopoda",
m e d i o - d o r s a lp a r t o l t h e m a n t l e c a v i t y e x t e n d s
Vol. 2, Plates25, 33, 37!) and which may degenerate
backwards between the stellate ganglia and fbnns a
during post-embryonic developrnentin certain forms
wide pouch, which forms a secondary, posterior
(Argonautidae). In other forms (Octopodidae) the
communication with the ventral mantle cavity in the
iateral parts are preservedas "cartilaginousrodlets"
areaof the gills,
in which the longitudinal axis of the
embeddedin the muscularmantie.Thus they remain as
gill is representedby a wide canal lying between the
supportingstructuresfor the mantle, indeedin the very
afferent and efferent vessels. in which the muscular
position where the tunnel retractor is typically
rnantleis connectedto the body by a powerful mid-
a n c h o r e d i n a l l t h e d i b r a n c h i a t e s( F i g . 6 4 ) . T h e
ventral muscle extending across the mantle cavity, (281) - in which the lemale sexualducts are bilaterally
observedin the Cirroteuthoidea,where its median part
symmetrical,the male duct is presentonly on the left
is preservedwhile the lateral, enlarged ends serve as
side,
in which the inner shell is extremely
strongestdevelopmentof the shell rudiment is
fin supports (cf. p. 34). No fossil records ol this
rudimentary, without anv hint of the typical
structureare available.Since it is so strongiy reduced
subdivisions.
and allows no detailed comparison with known decapodshells,a fossil octopodshell would probably
In additionto this generaldiagnosis.I give thoseof the
be
two extantsuborders:
nevertheless, one palaeontologist (Voltz) has
misidentified by
most palaeontologists;
gi n g 1 . C i r r o t e u t h o i d eaar e o c t o p o d sp o s s c s s i n u
discoveredit (p. 190). There is no reliable evidenceof
like, muscular fins set rvidelr apart on thc lateral
a conus, still less of any chamber formation. The
r66
b.
1\, l1-.
((ft')
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t' ty1l,i' L.--
\.*_i;j"
Fig. 97. -Reconstruction of Palaeoctopusnetvboldi from the Cretaceousof Syria (b). a. Eledonecirrosa (.Lam.).Young male from Naples,ventralview. .r,y, z: pa1lsof the hectocotylus. b. Perfectlyrecognizablebody of Palaeoctopuson a limestoneslab from Sahel-Alma,after Woodward, 1896,Pl. 6 (1/2nat. size).- k: beak,c: head.lr: funnel, lD: ink sac,f, fin. c. Octopttsdeflippii Vdrany from Naples.Ventral view and typical attitudeson the seabottom. Left: "stilting". Right: lurking behind branchesof a bryozoan colony, which it strikingly resembles.Note the eyes! Suckersin two rows 11/qnat. size). The difference comparedwith theseliving types rs Ihat Palaeoctopushas a relatively small head, a thicker mantle pouch, fins, and shorter arms. d. Octopus macropls Risso from Naples; juvenile form. (r/2 nat. size).
Short arms are observed in young animals of living
octopodids.The speciesshownherehas impressivelylong armsat the adult stage.(SeealsoFig. 98).
Cirroteuthoideaclosestto the type (probably the most
C. The genusPalaeoctopusWoodward 1896.
ancientones)are the Vampyroteuthidae,which retain a slightly cup-shapedshell in the dorsal part of the posterior end of the mantle. But there is no clear
Only one specimenis known, namely Calai'snewboldi de Sowerby 1846 -- Palaeoctopusnewboldi (de
distinction between the pro-ostracum and the conus
Sowerby)Woodward 1896 (Qu. J. Geol. Soc.,p.229)
(MadokaSasaki,1920,Proc.Un. St. Mus., Vol. 58, p.
(first cited as Calai's newboldi also by Woodward
zJ l.
{1896, Geol. Mag., p. 567}, a name that was subsequentlyabandonedfor nomenclatural reasons).
The extant Octopoda form two natural suborders, for which I have proposedthe namesPolypodoideaand
The animal is superblypreservedin a slab of limestone
Cirroteuthoidea(Naef, 1921).The oldest fossil genus
from the Upper Cretaceousof the Lebanon,presenting
Palaeoctopusis not easily accomodatedin one of these
a faithful picture of a true octopod without any further
suborders;it indeedrequiresthe creationof a suborder
reconstruction.Figure 97b shows only the undeniably
(Naef, 1921),which is of its own, the Palaeoctopoda
recognizableparts of the (286) fossil, without any
related to the extant groups in a similar way to the
restoration. This figure may be compared with the
relationship between the Belemnoidea,on the one
reconstructionby Bollo (1912, p. 126; see also Abel,
hand,and the Teuthoideaand Sepioidea,on the other.
1916,p. 83), which seemsto be not very successful.
t67 The animal looks particularly similar to a young O.
formatior.r of a typical shell inconceivable.
macroplts (Fig. 97d). The single-filearrangementof
Much more advancedjuvenile stagesof both sexes
the suckersis similar to extant speciesof Eledone (a).
(Fig. 98) also show the overallaspectof other octopod
The presenceof true fins marks a difference from all
larvae, along with a striking differencebetweenmales
polypodoids,the strikingly small head is a difference
and females,but there is no trace of any shell in the
from all extant octopods.The fins might be taken to
latter. The shell appearsonly at a certain body size
indicatethat this animal belongsto the Cirroteuthoidea.
when the dorsal arms grolr' markedly (288) longer than
But the special form and position of the fins, and the
the other arms and form a sort of loop. Within this loop
overall aspect of the animal argue against this
a skin fold spreadsto fonn the rudiment of the "shell
affiliation. Hence we should probably consider
membrane".Along the distal part of the arm, peculiarly
Palaeoctoptrsas a precursorof polypodoids,or as an
(289) enlargedskin glandsform, which actually secrete
intermediateform whose special position requires the
the "Argonautashell". I have unfortunatelybeen unable
creationof its own systematicunit at a (287) higher
to observethe very lirst stageof the forrnation of this
rank (p. 284). Unfortunately the shell rudin-rentis r.rot
rudiment; however, it can be reconstructedfrom the
visible. It probably had the character typical of
subsequentstructure and rnode of formation: the two
octopods as shown in Figure 4c, i.e. a transverse,
shell arms are bent backwards to cover the posterior'
archedplate providing a suppolt for the posterior end
end of the mantle pouch, using the latter as a mould for
of the strikingly inflated mantle pouch, and carrying
the secretion(by the two arms) of a cap-shapedinitial
the lateralfins.
s h e l l w h i c h r a p i d l y h a r d e n s .T h i s f i r s t r u d i m e n t afterwardsgrows in size in the way demonstratedby
D. The genusArgonuutaL.
more advancedstages:the glandularpart of the arm is applied to the free edge and forms a groove enclosing
Whereasthe lossil Palaeoctopusshowsno particularly
that edge,one row ol suckersbecoming attachedto the
striking features other than its intern-rediateposition
inside of the shell. The accornpanyingrnarginof the
between the recent suborders,the second octopod
skin bearssmaller glandularcomplexesthat secretethe
genuspreservedfossil is one of the most intrigr-ringand
inner (thinner) shell layer, whereas the otherwrse
interestinganimal types known. It has generateda large
similar, but stronger outer layer is formed by the
number of more or less justified interpretationsand
broaderglandularstrip of shell-formingskin (Fig. 98d).
hypotheses,in contrastreasonableconsiderationand
Early juvenile shells have been figured by Hoyle
observationhave been largely neglected.A detailed
(1904, Pl. 10, Fig. l2): but their earlier structurecan
description of the anatomy and development will
also be recognizedfrorn older shells by following the
appearin my monographof the living cephalopods;in
growth stagesbackwards. Tire following stagesthen
the sectionsalreadypublished(Vol. II, Pl. 32-37),the
appear:the early cap- or bowl-shapedrudiment (which
embryonic stages are comprehensively described,
doesnot show much evidenceof its bipartiteorigin. i.e.
startingfrom the cleavagestages.They are very similar
the paired shell arms) first grows by the concentric
to other octopod stages and, like them, show the
addition of material (Fig. 98d). Only subsequentlywill
formation of a small shell sac as the internal shell
the ventral margin grow faster, whereas the dorsal
rudiment. In this form, however, the shell sac rs
margin grows out laterally and forms the increasingly
particularly underdeveloped,composedof very few
thicker "columella".The border forms a sharp angle,
cells, and barely visible at the end of embryonic development.- The hatchling is a true, small octopod
whose growth is often acceleratedand thus forms
and closely resemblesthe correspondingstagesofother argonautids(loc. cit., Pl. 3l). Thereis not the slightest
"ears" like certain ammonite shells: these structures r n a ys u b s e q u e n tdl ye c r e a sien s i z e .The shell, which at later stagesforms ribs, keels
trace of any externalshell. The "mantle" (which is nol
and peripheraltubercles,housesthe rapidly growing
prirnary mantle of
female; she holds the shell with her reflexed arms and
tetrabranchiates, as mentioned on p. 22) bears the
swims in an attitudesimilar to the defensiveposition of
integumentarysetaetypical of octopod larvae and thus
an octopodsitting in a den (Figs 98 and 100).The eyes,
makes secretoryactivity that could be relatedto the
mouth and beaks, together with the strong arm bases
homologous with
the
168 Fig. 98. Morphologyof the genusArgonauta. a. Young female of A. argo L. from a plankton sample taken at Naples, without an external shell, after total
,wfu
reduction of the inner shell rudiment (which was present during
embryonic development), and before the
transformationof the dorsal arms. What is shown here is a typical young octopod, very similar to the corresponding : stagesof Octoprlsspecies1/r.i t. 1 nat.size. b. A corresponding stage of the male, from the same samples.The mantle cavity has openedup due to shrinkage of the muscular mantle, the press-button connection between the mantle and funnel attachmentsis interrupted, so that the inner parls of the funnel complex are visible. The left arm of the third arm pair begins to be modified as a hectocotylus.5/1nat. size. c. A dorsal arm of the female shown in a: it is similar to the other arms, bearing the three 1arva1suckers(1-3) plus the rudiments of four more suckers which will subsequently multiply like leaf buds on the vegetative cone of a 25 cormophlrestem. 1 na1.size.
*.1:i:-:
c/. A young female of the same species,with fully formed shell arms and a rudimentaryboat-shapedshell. 3: growing 1: the shell margin, .2:upper margin forming the co1umel1a, auricular corners of the latter, the growth of which will subsequentlybe acceleratedor decelerated,4: shell rib, 5: protuberanceon the keel. 3 l2nat. size. e: dorsal arm of an adult female. "creeping" (by rneansof
,4
the advancing,elongate suckers)from the shell cavity. frnally to cover the outer shell surfaceentirely. 12: sucker
stem, B: intermediatcmembrane,2: arm axis, 9: columella, ,10:growing margin, 1J: rib, l4: ventraltubercle, 11: black substance, l/a secondarilyaddedto the upperpart ofthe venter. nat. size. / Totally expandedright dorsal arm, loop-shaped,with extendedshell membrane.1: inner skin fold connectingthe suckers.7: outer skin fold,2: free edgeof shell membrane,3:junction betweenits initial and terminal(4) parts,5: part of the arm enclosingthe growing margin of the shell, both inside and outside,accompaniedby a glandularstrip which adds material to the shell margin (6), indicatedhere only by the shell gland tubules. g. Maturc female, r/a nat. size (same individual) shown inside its shell in typical swimming position. The left dorsal arm is represented as ifcut, the shell transparent. ft: tips ofmandibles,e: eggs,in all developmentalstages,r: columella. /zand l. Young Octopusvulgaris Lam. from the planktonat Naples,essentiallylike a. 5 I nat. size. t. Young O. vulgari,safter changing to a benthic life style; except for the normal dorsal arms, the animal is similar to that shown in d. 2/1 nat. size. C/r: chromatophores,Si: velar membrane,Plr: pupil, O/: upper 1id,l/: outer lid, Ro: olfactory organ,Tt: funnel pouch, Ir: funnel tube"Hh: skin tubercle.
with their suckers,are ready to act while the rest of the
Given these facts, there is no basis for any attempt
animal is hidden inside the shell. (Such an attitude
to find homology between this peculiar apparatusand
generally remains visible in a preserved speclmen
an ammonite shell, as will be realizedby anyonewho
without its shell, indeedeven in other octopods,as can
may still have been under a different impressron.
be seenin museum coliections).When hard-pressed,
Nevertheless,the overall similarity betweenthese two
the animal can leave its shelter, returning to it later,
structuresis very striking; a tentativeinterpretationwill
(290) in strict contrast to the normal relationship
be given below (p. 292). A peculiar sort of "allusion"
betweena mollusc and its shell.
to the well known nautilus shells must also be
169 mentioned. The earliest part of the argonaut shell, which progressivelypoints upwards and forwards,is s u b s e q u e n t l yd e c o r a t e dw i t h a " b l a c k s u b s t a n c e " secondarilyaddedliom the outside;this formation rnay have an ecological interpretation:this part of the shell often remainsuncoveredby the dorsal arms and, if left as a light spot, would spoil the protectivecolorationof the rest ol the animal. (The differentiationof the shell tneubrane which covers the whole structureduring slorv su'imming {Fig. 100} indeed provides the possibility of integration with the process of chromatophoric color change. * For the different
Fig. 99.
Fossillrgonarrla she1l.A. sisrttondaeBellardi,
1872,from the Plioceneofnorthem Italy (3/anat. size).After v. Stromer(1909).
interpretationof the black substancein Nautilus see "Cephalopoda", Vol. I, p. 62').
form; for in the life of the animal they assumethe same
The young fernaleArgonauta startsto attacheggs
function as an egg carrier (cf. "Cephalopoda", Vol. I,
insidethe apex olthe shell at a surprisinglyearly stage,
chapter54). The closestrelative of Argonauta, Oc.vthoe
thus demonstratingits dominant biological role as a
tuberculata Raf. (which can be placed in the same
brooding device, as alreadysuggestedby the exclusive
subfamily), has become completely ovoviviparous
occurrenceof shells in females. Under natural
(evenArgonatta depositsthe eggs only after cleavage
conditionsit is likely that the newly hatchedyoung find
and formation of the germinal layer!); Ocythoethus no
some temporary protection inside this brood chamber.
longer needs a brooding apparatus.However, in the
But given their typically planktonicnature,they will
male Ocythoe, we observe a peculiar instinct: the
not actually need post-embryonicbrooding. Their very
animal adoptsan empty Salpa barrel, or emptiesa full
small size (1.4 mm) protects them fairly well from
one, and adoptsthe same position as an Argonanta
many predators,along with their large number (several
female inside its brood shell; the male of Ocythoeth:us
hundredthousandfrom one mother).
drifts about as a pelagicDiogenes(cf. Jatta,1896,Pl. 7,
The above description, which is merely a
Fig. 8). Even the female Ocythoe,when captive,tends
preliminary note, is given here to refute the repeatedly
to exhibit the same attitude as a female Argonttuta tn
expressedview that Argonauta were deriveddirectly
her brood shell. Most preservedspecimensshow this
from ammonites, and to permit an objective
pattern of reflexed arms. The dorsal arrn loop has a
understanding of
membrane reminiscent of the shell membrane of
the
fossil
relatives (cf.
Vol. 1, chapter56). "Cephalopoda", The brood shells of females are indeed the sole fossil remains available from extinct species.They
Argonauta, but it is devoid of the specialglandular complex. Based on these facts, I propose the following
occur fiom the Miocene onwards and are similar in
hypothesis: (292) The ancestor of the extant
every respectto the extant (291) forms, without being
argonautidsadoptedempty shells at the adult stageto
totally identical.Thus we learn nothing from the fossil
lay their eggs in. The eggs were fixed to the inner
brood cases about simpler preliminary stages of
surfaceof the shells (as in living octopus)using the
morphogenesisq6.
secretionsof the skin glands. Subsequentlythese
Indications of the phylogeneticorigin of the brood
secretions were also usedto enlargethe brood case,in
sheil of lrgonaltta rnay be derived from the following
a similar way to whal Adantsia palliata achievesin
facts: a diff-erent, extant argonautid, TrentoctopLts
adding materialto a gastropodshell for the hermit crab
t.,iolaceusD. Ch., produces(fiom secretionsof the
inhabitingit. Finally the foreign shell servedonly as
glandularcomplex of the dorsal arms) rodlet-shaped
the nucleusof the shell producedby the cephalopod
bodies to which the eggs are attachedand thus carried
and ultimatelybecamedispensible. This hypothesis
b y t h e f e m a l e . T h e s e b o d i e s c a n b e c o n s i d e r e da
provides an answerto the following question:how has
hornologrie of
the brood shell of Argonauta,
such a perfectly formed structureappeared"suddenly"
notrvithstanding the very limited similarity of overall
within an otherwisehomogeneous group?Biologically
170
Fig. 100.
Argonauta argo L. (tl2 nat. size). From live observationsin the aquarium, drawn post mortem y'oz sketchesand
photographs(From "Cephalopoda",Vol. l, P1.1.1). Above: Adult female with dorsal arms fu11yspreadout over the shell. Below: The same animal, with dorsal arms withdrawn, another typical attitude which is rapidly assumedin the presenceof obstructions.The dorsal arms with their shell membraneare stowed away inside the shell exactly like the other arms. Inset:The adult dwarf ma1e.which is devoid of a shell.drawn to scale.
t71 it doesnot make senseto expectthe formation of such
(ribs, keel, peripheraltubercles)by Argoncttttais due to
an apparatuswithout any relation to a more easily
some kind of transf-erof plastic sense as it u'ere
achievable structllre. That foreign shells may be
(personallyI ar.nconvincedof this, and I hope to retlll'n
adoptedfor brooding pulposesis der.r.ronstrated e.g. by
to this view elsewhere).
Phronima sedentaria,an amphipod, the female of
Fossil lrgonauta
shellsare rarely preserved;
which uses a Salpa barrel to keep the eggs lvith her
similarly the brood shellsof the extant "papernautilus"
when drifting in midwater. Another instructive
are rarely found intact without the animal. Fragments
exarrple,more closely relatedto Argonauta,is Octctpus
sometimesoccur in dredgesamples.Thus the fact that
digueti, which uses empty bivalve shells as brood c h a n . r b e r s( c f . P r o m e t h e r i s , V o l . g , 1 8 q 7 . o r
we lack an uninterrupted,reliable fossil record of the
Rochebrune,1896, Nouv. Arch. Mus. Hist. N. Paris,
n o ta r g u ea g a i n sot u r a s s u r n p t i o n s .
Vol. 8,p. 75).
genus(p. 291) extendingback to the Cretaceousdoes The oldestknown Argonauta is A. johanneusHilber
Moreover, my assumptionrnay possibly explain the
( 1 9 1 5 ,p . 1 0 7 ,P l . 1 , F i g . 1 - 2 ) .I t w a s f o u n d i n t h e
peculiar sirnilarity which exists betweenArgonaLtta
Miocer.reof the Steiermark.Another fossii specresrs
a n d c e r t a i n U p p e r C r e t a c e o u sa m m o n i t e s : i t i s
known liom Japan: l.
conceivable that the gas-filled, buoyant shells of ammoniteswere particuiarly useful as potential brood
),oshiv,araen. sp. (cf. Y o s h i n ' a r a ,1 9 0 1 . p . 1 1 4 , P l . 5 ) . I t w a s f o u n d i n
chambersfor the ancestorof our extant argonautids;
Neogene blue-grey tuff without a precise age d e t e r m i n a t i o n .- A f o s s i l l r g o n a u t a s h e l l r . v a s
this assumptionindeed forms a complementarypart of
describedby v. Eichrvaldin 1830 (.A.zborzev,ska)who
our hypothesis.I thr"rsassumethat the protoargonautids
c o n s i d e r e di t t o b e a l b r a r n i n i f e r a n ,a n o t h e r b y
adopted empty arnmonite shells and subsequently
Boenninghaus(A. c'ornu) ivho classifled it as an
becameadaptedto them in the way describedabove.
ammonite.A. ,sismondue Bellaldi (Liguria,p. 11,Pl. l.
Sincethe octopods(Polypodoidea)show a tendency
Fig. l; or.rrFig. 99 herein) rvas describedfi'orl the
to crawl into any availablecrevice and to settlethere''-,
Plioceneof Piedmont,along rvith ,4. fizan.sSolander
it seemsmore than likely that the extremelynumerous
1 7 8 6 ,a n e x t a n ts p e c i e sS. e ea l s oB t i l o w ( 1 9 2 0 ,p . 2 1 9 )
empty ammonite shells (293) occurring along coast
a n dB e l l a r d (i i 8 3 8 ) .
lines were quite often adopted by these octopods. Argonauta, in its turn, succeededin becoming (291) independentof these foreign shelters;this must have happenedat the end of the Mesozoicera, i.e. during the Upper Cretaceous,since ammonitesbecame extinct and thus were no longer available,while nautilid shells were too uncommon to provide an equivalent substitute,apart from the fact that their shapewas less suitable(the shoft living chamber{p. 20} of a drifting shellhangsdownwards). We must of course assume that the whole argonautidfamily has first evolved in the suggested direction.But only rn Argonattta has the result been fuily conserved. Oc,,-tho€and Tremoctoptrs apparently had not yet reacheda similar degreeof perfection and had to find other solutions vn,henthe ammonites became extinct (p. 29i): in other words they were forced to do without the extraneousabode,not being ableto producea completesubstitutethernselves. The interestingquestionariscshere (without being pursuedfurther) as to whetherthe apparentimitation of the shell ornamentof certain Cretaceclus ammonites
=
172 (295)
apart from the effects of extraneous obstacles and
Conclusion.
personalfailure. But we believe that we have pointed
Throughout the preceding discussions,ranging over a
have improved by sheddingnew light on it. -
out a way which is not new, but which we think we
very wide field, we had two aims in mind, one special
To conclude we provide an overview of the
and one general.One aim was to find and describe
completeclassificationof living and fossil dibranchiate
natural order in a hitherto confusing diversity.
genera, which we regard as the framework of a
Experiencedreaderswill recognizethat we have been
systematicsynthesis.
successfulto a cefiain extent,especiallysincewe draw special attention to the remaining gaps 1n our knowledge. Future researchwill probably add new
Families and genera of dibranchiate cephalopods.
elementsto the presentsketchso as to improve it. Will it ever be possibleto "clarify" the historical courseof
First order:DecapodaLeach 1818.
evolution by progressivelycombining closely related speciesin truly phylogenetic sequences?I have some
Subordera) t BelemnoideaNaef 1912.
doubts and at best dare to hope that it may be possible for the belemnitefamily. If we have neverthelessbeen
l . A u l a c o c e r a t i d a eB e r n a r d 1 8 9 5 . A u l a c o c e r a s
able to obtain a full picture from innumerabledetails,it
Hauer 1860, Dictyoconites Mojs. 1902,
was due not to mere compilationof many partsbut to a
Calliconites Gemm. 7904,Atractiles Giimb.
systematicsynthesis.
l 8 6t .
This leads us to our second aim: we wish to demonstratewith this monograph how palaeontology can work methodicallyas a biological discipline, in other words we wanted to provide a model of true "palaeobiology"in which the principlesof this science
2. Phragmoteuthidae Naef 1921.Phragmoteuthis Mojs.1882. 3. XiphoteuthidaeNaef 1921.XiphoteuthisHuxley 1864. 4.Belemnitidaed'Orb. 1845. (a) Hastitinaenov.:
can be explained.It was essentialfirst to define the task
Hastites Mayer 1883, Rhabdobeltrsn. g.; (b)
and the generalconditionsunder which an answer can
Coeloteuthinae nov.: CoeloteuthisLiss. 1915;
(if possible)be given, before drawing the logically
( c ) P a s s a l o t e u t h i n aneo v . ' . N a n n o b e l u s
m o s la c c e p t a b cl eo n c l u s i o n s .
P a w l o w 1 9 1 3 , P a s s a l o t e u t h i sL i s s . 1 9 1 5 ,
We based our work on the idea that the observed
Pseudohastitesn. 9., BrachybelusNaef 1922,
diversitl; offorms is the (296) expression oJ'a process
Homaloteurh ls Stolley 1919, Megateuthis
advancing throtrgh time, namely a modi/ication of
Bayle 1878,Gastrobelusn. g., Pleurobelusn.
morphological norms. The ultimate objects of our scientific interest are these very norms (rather than
9., Odontobelus n. g., SalpingoteuthisLiss. 1 9 1 5 , D a c t y l o t e u t h l sB a y l e 1 8 7 8 ; ( d )
h y p o t h e t i c a l g e n e t i c r e l a t i o n s h i p s ) .T h e s e n o r m s
Cylindroteuthinae ( 2 9 7) nov.: Cylindrotetrthis
indeed are prerequisitesfor the presentationof the
Bayle 1878, Oryteuthis Stolley l9l l,
observedfacts, and the presentationin its tum is a way
AulacoteuthisStolley 1911,Rhaphibelusn. g.;
of testing the preliminary assumptions.The types
(e) Belemnopsinaenov.: Belemnopsls Bayle
representedthe basis for the reconstructionof fossil
1818,Hibolites Mayer 7883,Belemnoconus11.
forms, and the resulting pictures (close to nature and
g., Parahiboilres Stolley 1971,Mesohibolites
taking account of special conditions) in turn provided
Stolley l9ll,
confirmationof the initial concepts.That at least is the
Belemnitellad'Orb. 1845,ActinocamaxMiller
N e o h i b o l i t e s S t o l l e y 1 9 11 ,
situation now that we have learnedto view the whole
7823, Dicoeiiles Bcihm 1906; (fl Duvaliinae
field.
(Pavl.): Duvalia Bayle 1878,Pseudoduvalia
It is evident that this aim can only be approached step by step given the nature of the undertaking.lt is
n. 9., PseudobelusBIarnv. 1825,Conobelus Stolley 1919,RhopaloteuthisLiss. 1915;(g)
also evident that total perfection cannot be hoped for,
Bayanoteuthinaenov.: Ba1;anoteuthlsMun.-
=
173 Ch., Stvracoteuthis Crick. Zitt. 1885) Naef 1921. 5 . B e l e r n n o t e u t h i d a( e Belemnoteuthls PearceI 842. 6.Diploconidaenov. fam. Diploconus Zrtt. 7868. Conotetrthis d'Orb. 1842,Amblvbelusn. g. 7.VasseuriidaeNaef 1921. Va.ssetirla Mun.-Ch. I 880.
(2e8) 6) Metateuthoidea oegopsida ( d ' O r b .1 8 4 5 )N a e f 1 9 2 1 . 1.BathyteuthidaePfeff. 1912.Bathvteuthis Hoyle 1885,Ctenopler-rxAppellcif I 889. 2. GonatidaeHoyle 1886.BerryteutlrlsNaef 1921, GonatusGray 1849,Gonatopsi.s Sasaki1920. 3 . O n y c h o t e u t h i d aG e ray 1849. Onvchoteuthis Lichtenstein1818,Ancistroteuthi,s Gray 1849,
Suborderb) TeuthoideaNaef 1916.
T e l e o t e u t h i . sV e r r . 1 8 8 5 ,O n v c h l a L e s u e u r 1 8 2 1, C h a u n o t e u t h i sA p p e l l o f
a) 'i' PrototeuthoideaNaef 192I . l . P l e s i o t e u t h i d aNea e f 1 9 2 1 .P a r a p l e s i o t e u t h i s Naef I 921, Plesioteuthis Wagner 1860, Sl'loteurhisFritsch I 9 I 0.
l89l,
Tetront'choteuthisPfeff. 1900,Moroteuthis V e r r . 1 8 8 1, C y c l o t e u t h l .Ji o u b i n l 9 l 9 e t ( c f . Cephalopoda, vol. I, p. 48). 4. NeoteuthidaeNaef 1921. NeoteullzsNaef 1921.
2. LeptoteuthidaeNaef 1921. LeptoteuthisH. v. M. l 834.
5. OctopodoteuthidaeBeny 1912. Octopodoteuthi.s Riipp. 1844,Cuciotettthis Steenstr. 1882.
3 . G e o t e u t h i d aN e a e f 1 9 2 1. G e o t e l t l r l s M i l n s t . 18 4 3 . ,1.BelopeltidaeNaef 1921. BelopelllsVoltz 1840, P a r a b e l o p e l l l . s N a e f 1 9 2 1 ,L o l i g i n i t e s (Quenst.1849)Naef I 921. 5. Lioteuthidaenov. fam. Lioteuthisn. g.
6.HistioteuthidaeVerr. 1881. Histioteuthis d'Orb. 1839, Calliteuthis Yerr. I 880 (Sllgratctteuthis Pfeff. 1900), Histiopsi.s Hoyle 1885. Meleagroteutftrs Pfeff. 1900. 7 .A r c h i t e u t h i d a e P f e f f .
1900. Architeuthis
(Steenstr. 1857)Verr. 1880. 8. Enoploteuthidae Chun 1910. (a) Pyroteuthinae:
Naef 1921. B) t Mesoteuthoidea 1.TrachyteuthidaeNaef 1921. Trachl,teuthisH. v. M. 1846,Qlyphiteuthis Reuss1870. 2.Beloteuthidae N a e f 1 9 2l . B e l o t e u t h i sM t i n s t . I 843.
P ) ' r o t e t r t h i s H o y l e 1 9 0 4 ,P t e r v g i o t e u t h i s Fischer 1896; (6) Lycoteuthinae'. Lycoteuthis P f e f f . 1 9 0 0 ,L a m p a d i o t e u t h i s B e r r y 1 9 1 6 , Nematolontpas
Berry 1913; (")
Enoploteuthinae'. Enoploteuthi,c d' Orb. I 839.
3. PalaeololiginidaeNaef 1921. Palaeolo/lgo Nael
Abralia Gray 1849,Abraliopsls Joubin 1896,
1921(:TeuthopslsWagn. 1860),Tusoteuthis
Ancistrochit"ersGray 1849, Thelid ioteuthis
Logan 1898" Ph,"lloteuthisMeek & Hayden
Pfeff.1900.
1860(?).Ptiloreurhis Gabb 1869(?). 4.Celaenidae N a e f i 9 2 l . C e l a e n oM i i n s t . 1 8 9 2 , Celuenoteuthis n. g.
9. Psychroteuthidae Thiele 1921. Psvchroteuthis Th.1921. 1 0 . O m m a t o s t r e p h i d aG e i l 1 . 1 8 7 1 . 1 1 l e -S r teenstr. 1880, Todaropsls Girard 1889, Nototodarus
y) Metateuthoidea myopsida
P f e f f . 1 9 1 2 ,O m m a t o s t r e p f t ed. s' O r b . 1 8 3 5 ,
( d ' O r b I 8 4 5 )N a e f 1 9 2 1.
Dosidicus Steenstr.1851, Ht,aloteuthisGray
1 .L o l i g i n i d a e S t e e n s t r .1 8 6 1 ( a s " L o l i g i n e i " ) .
1849, Sthenoteuthis
Verr. 1880,
Loligo Lam. 1799,SepioteuthisBlainv. 1924,
S!,mp I ec t o teut h i s Pfeff . 7900, E uc I eo t eut h i.s
L o l i c t l u s S t e e n s t r .1 8 5 6 ,D o r y t e a t i l s N a e f
Beny 1916.
1912.Alloteurlrls(Naef MS) Wrilker 1920. 2. Promachoteuthidae Naef 1912.Promachoteuthis
J o u b i n1 8 9 5 .
Troschel1857. 12. BrachioteuthidaePfeff. 1908. Brachictteuthis
Hoyle1885. 3 .L e p i d o t e u t h i d a e N a e f
11. Thysanoteuthidae Keferst. 1866. Thvsanoteuthis
1912. Lepidoteuthis
Ven. 1881. 13.ChiroteuthidaeGray 1849. (a) Chiroteuthinae: Chirotetrthisd'Orb. 1839,ChirothaumaChun
t74 1910; (b) Mastigoteuthinae MastigoteLtthis Verr. 1881 (Chiroteuthoides Berry 1920, IdioteLrthis
1881. 9. SepiolidaeKeferstein 1866. (a) Sepiadariinae
Sasaki 1916); (c)
N a e f 1 9 1 2 : S e p i a d a r i u mS t e e n s t r1. 8 8 1 ,
Grimalditeuthinae(GrimalditeuthidaePfeff.
Sepioloidead'Orb. 18a5; (b) RossiinaeNaef
1900):
1912: Ro.ssia Owen 1834,SemirossiaSteenstr.
Grintalditeuthis Joubin 1898
EnoptroteuthrsBerry 1920.(299)
1 8 8 1 ; ( c ) H e t e r o t e u t h i n a eN a e f 1 9 1 2 :
14. Cranchiidae Gray 1849. (a) Cranchiinae:
HeteroteuthisGray 1849,NectoteuthisYerr.
C r a n c h i aL e a c h1 8 1 7 L , e a c h i aL e s u e u r1 8 2 1 ,
1883, Iridoterrllrls Naef 1972, Stoloteuthis
Pyrgopsis Rochebr. 1884,Liocranchia Pfeff.
V e r r . 1 8 8 1 ; ( d ) S e p i o l i d a eN a e f 1 9 1 2 :
1 8 8 4 .L i g u r i e l / a I s s e l 1 9 0 8 ;( b ) T a o n i i n a e :
Sepiolina Naef 1912,Euprymna Steenstr.
Phasmatopsi.s Rochebr. 1884, ToxettmaChun
1 8 8 7 , S e p l o l a L e a c h 1 8 1 7 ,S e p i e t a N a e f
7906, Taonius Steenstr.1861,Desmoteuthis
1912,RondeletiolaNaef 1921.
Verr. 1882,Megaloc'ranchiaPfeff. 1884, Teonidium Pfeff. 1900, Chr,,-stalloteuthis Chun 1906,PhasmatotettthionPfeff. 1912,
(300)
Galitetrthi.s Joubin 1898, Corltnomma Chun
Secondorder:OctopodaLeach1818.
1906, B a tho tha uma Chun 1906, Ver r i I I i t etrth i s B e r r y 1 9 1 6 ,L e u c o c r a n c h i a J o u b i n 1 9 1 2 ,
Subordera) t PalaeoctodaNaef 1921.
Hansenioteutftzs Pfeff. 1900,Sandalop.rChun 1906, Helicoc'ranchia
M a s s y 1 9 0 1,
T e u t h o w e nC i ah u n I 9 I 0 .
1 .P a l a e o c t o p o d i d aDeo l l o 1 9 1 2 . P a l a e o c t o p u s Woodw. 1896.
1 5 . Jonbiniteuthidaenov. farn. p. 299.Joubiniteuthis Berry 1920ee.
Suborderb) Cirroteuthoidea Beny 1920. 1.Vampyroteuthidae Thiele 1915.Vampl;roteuthis
Suborderb) SepioideaNaef 1916.
Chun 1903, Watasella
Sasaki 1920,
M e l a n o t e u t f ti s J o u b i n 1 9 1 2 ,L a e t m o t e u t h i s 1. tBelemnosidaeNaef 1921. BelennoszsEdwards 1849,Belemnosellanov.,Spirtrlirostrella
2.CirroteuthidaeKeferstein 1866. Ciruotetrthis
Naef 1921.
E , s c h r i c h t1 8 3 6 , S t a u r o t e l t t h i sV e r r . 1 8 7 9 ,
2 . t B e l o p t e r i d a eN a e f 1 9 2 1 . B e l o p t e r a ( D e s h . ) B l a i n v . 1 8 2 5 ,B e l o p t e r e l l a N a e f
Beny 1913,Hymenotetrthls Thiele 1916.
1921,
Belopterina Mun.-Cg. 1872,Belopteridium n. 6.
Froekenia Hoyle 1908, Cirrothauma Chun 1917,Chuniotettthis Grimpe I9I6. 3.Opisthoteuthidae Verr. 1896. Opisthoteuthis V e m .1 8 8 3 .
3 . t B e l o s e p i e l l i d a eN a e f 1 9 2 1. B e l o s e p i e l l a Alessandrini1905. 4 . t S p i r u l i r o s t r i d a eN a e f 1 9 2 1. S p i r u l i r o s t r a
Suborderc) PolypodoideaNaef I 92 L
d'Orb. I 841,SpirulirostridiLtmn. g. 5. tSpirulirostrinidaeNaef 1921. Spirulirostrina Canavari1892. 6.Spirulidae (d'Orb. 1826) Owen 1836. Spirula L a m .1 8 0 1 . T . S e p i i d a eK e f e r s t e i n 1 8 6 6 . ( a ) f B e l o s e p i i n a e
o) Ctenoglossa Naef 192ltoo. 1.Amphitretidae Hoyle 1886. Amphitrettrs Hoyle 18 8 5 . 2. BolitaenidaeChun l9ll.
Bolitaena (Steenstr.
1859, Hoyle 1886) Chun 1904, Eledonella
Naef 1921 Belosepia Yoltz 1830; (b)
Verr.
SepiinaeNaef 1921: Sepia L. 1758,Sepiella
VitreledonellaJoubin 1918.
G r a y 1 8 4 9 , H e m i s e p i u s S t e e n s t .1 8 7 5 , MetasepiaHoyle 1885. 8. IdiosepiidaeAppellof 1898.Idioseplls Steenstr.
1884
(Japetella Hoyle 1885),
175 Naef I 921. B) Heteroglossa
( P o l a e o c t o p a s ) m u s t b e c o n s i d e r e di o s t , a n d t h a t
l . O c t o p o d i d a ed ' O r b 1 8 4 5 .O c t o p u . L i am. 1799,
representshundredsof types..Evenmore striking is the
E l e d o n e L e a c h 1 8 1 7 ,V e l o d o n aC h u n 1 9 1 5 ,
fact that we have no fossil speciesof the 17 families of
C i s t o p t r s G r a y 1 8 4 9 ,P i n n o c t o p u s d ' O r b .
metateuthoids,most of which must have originated in
I 845.
the Upper Cretaceous.Here again hur.rdreds of types
2. ArgonautidaeNaef 1912.(a) ArgonautinaeNael 7 9 2 7 :A r g o n a u t a L .
1758, Ocythoti
have been destroyed.The most curious fact is the
-
absenceof sepioid decapods(with caicilied shellsl)
Rafinesquel8l4: (b) Tremoctopodinae Naef
fiom the Lower Lias to the Eocene,althoughwe are
1921.:TremoctopusD.-Ch. 1829,Alloposus
forced to adrnit their special relationship to the
Ver. 1880.
teuthoidsas opposedto the belemnoids(p. 25. 167.
: :
189). Since the phragmoconewas alreadylacking in the prototeuthoidsfiom the Lias e fl-ower Toarcian],a This lirst (graphical)overview, which does not take
c o m m o n a n c e s t o r c a n o n l y b e e n v i s a g e di n t h e
account of temporal relationships,is followed by a second overvierv fitting the morphological (301)
lowermostLias. at the latest.
diversity into the systemof geologicalformations.This
branchinglineages(or merginglir.reages when viewed
combination of typical relationshipsand stratigraphic
from the present). Therefore the assumption is
data indeed permits a sort of historicalpresentation.
inevitable that most speciesbelonging to the stem
The orderly presentationin a sort of genealogicaltree
lineagesare as yet unknown. On the basis of such
rests chiefly on systematicmorphology (Naef i910,
negativeevidenceit is of course impossibieto drarv
p.20),much less or.rpalaeontological sequences. There
generalconclusionsas to the true courseof evolution.
is no claim to consistentlyshow all the relationsacross
(Most of the gapscanbe explainedby the conditionsol
the geologicalformations.There are enonnoLrsgaps in
preservationwhich u''erevery unfavourable fbr most
the fossil record.We presentonly the most essential
the sepioids species:the octopodshaveno solid she1ls,
points:
started out with extremely small littoral fonns). This negativeresult,on the otherhand,justifiesa separation
Among the living octopod families only the
\\,e have no continuous. Even for the belernr.rites
Argonautidaeinclude fossil representatives. Since it is
of (even historical) morphological (302) from
impossibleto link the whole order to known Mesozoic
phylogeneticstudies,which must be limited to the
or Cenozoic forms (a11of which are specialized
cases u'here their specific conclusions can be
decapods), all predecessors and extinctrepresentatives,
supported,namely by testing the criteria of blood
with the sole exception of a problematic fossil
relationship.
-
=
=
t76
Fig. 101. Phylogencticoverview ofthe dibranchiateccphalopods.A tree-likediagramis relatedto the stratigraphicsystems.This is indispcnsablefor visualizingsystematicrclationshipseven ifone doesnot envisageactualgcneticaffinities. It expresseswhat we can safely assumeabout the evolution ofthc group. The "x" marks indicatethe positionsofassumed evolutionaryseriesor evolutionary transfbrmationsof types, points where actual fossils can be accommodatedin terms of time and morphology.
They reveal the
stagesat which the modifications of certain types had arrived at a given time, whcrcas the phasesof other metamorphosesare unknown.For example.we do not know how the "protosepioids"really looked at the Middle Jurassic,Upper Jurassicand Cretaceous levels. and whether they already conformed in all details to this designationin the senseof'the definition given on p. 38. It is perfectly conceivable that they still had belemnoid shells, and that the modification of thc she11only occurred in the Upper Crctaceous(Diploconidae?cf. p. 3 i).
177
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179 Notes
prof-essional palaeobiologistis rnore knowledgeable than a zoologist about animal ecology, whether on
') SeeA. Naef, Die Cephalopoden, Monograph35 in:
special or generalaspects.It should certainly be
Fauna and Flora of the Gulf of Naples. 1st issue
r e c o g n i z e d t h a t A b e l ' s " P a l a e o b i o l o g yo f t h e
with 56 platesaccompanyingvolumesI and II. R.
Cephalopoda"(1916) containsuseful elementsin
Friedlrinder, Berlin
the
1921 (here cited as
"Cephalopoda").
preliminary
study
of
extant forms,
notwithstandingthe inclusion of unintentionally misleadingdata from the literature.But this part of
2) This doesnot deal with the descentof individuals!
the work is in lact concernedwtth zoolog.v.On the other hand, it must be said that Abel's study lacks
'r) Disregardingspecialmorphologicalconceptsdealing
the systematicmorphoiogical foundation that is
with partial phenomenain natural beings (gastrula,
needed; the latter can only, be provided by a
pteryglum.etc.).
specialisthaving a long experienceof indir,idual research.
a) What is characteristic,essential,typical is apparently cletermined by the hypothetical agents that are inheritedlrom one generationto the next as a stable
o)
That Steinmann'svieu, can apparentlybe neither refuted nor confirmed is due to the poor basis of
totality of "all hereditaryfactors" or "genotype".
both his own view and that of his opponents.ln
This does not itself have an uninterrupted existence!- The germ cells contain it only as a
replacing fancifiil
potential,not as a manifestfbrm. and there is no
deliberately demand acceprance of
way ol comprehendingand describingit directly as
i n d i s p e n s a b l e b, a s i c r u l e s o l c o m p a r i s o n a n d
a really effectualthing inside thesecells. Therefore
syntheticviewing.
phyiogenetics with
a
methodically sound systematicmorphology, we the
t h e v i s u a l i n - r a g e( p h e n o t y p e u n d e r c e r t a i n conditions)is used as a syrnbolic representationof
The suggestiob n y S . T s c h u l o k( 1 9 1 0 )t o c o n s i d e r
the genotype"even in the scientific study of the p h e n o m e n ao f h e r e d i t y . " G e n e s " c a n o n l y b e
"genetics",in the senseof evolutionarytl.reory.as a specialbiologicaldiscipline,to be placedalongside
characterizedand localizedby their effects. Thus
systematicsand morphology,is out of the questior.r,
a n o n - p e r c e p t u apl o s t u l a t e o f c a u s a i t h i n k i n g
indeed surprisinggiven the compactnessof most of
provides the only thorough conception of the
Tschulok'sideas.Such an innovationwould mean
continuity of life, which is certainly perceptr-ral as
picking the fruit from the tree of sciencebefbre it is ripe (cf. Tschulok,1922,p.2).
"parental reproduction", yet is not r.eally understandable in scientificterms (cf. Naef-.1919. p . 38 - 4 1 ) . O n l y t h e f o r m a t i v e t y p e [ G e r . : Bildungsnorrn]as such appearsdurable.Given this
" ) The "cicatrix", lor example,can be called a shell nucleus,this structurecan be found on the outer
limitation. even in lineageswhich can actually be
surfaceof the apex in variousNautiloidea,and has
observed,it is certainly necessaryto ernploy a morphological-systematic approachto forms whose
often been interpretedin fanciful ways (cL e.g. Banande,1877).
blood relations cannot be recognized(discarding any assumptionaboutpossibleblood relationships), rather than to approach them from a pseudogenealogicalperspective.
, ) As a curiosityone may quoteRiefstahl(1886)who contendsthat: '"Thephragmocone of the belemnites grows by intusception.""The secondaryseparation of the septa results liom the growth of the
s) The same is true, of course,of the assessment ol Iheir mocleo/ lije. In this respect,O. Abel (1921, p.
interveningzones(betweenthe septalinsertions)of t h ep h r a g r n o c o nuea l l " .
134) makes a rather strange suggestionby proposing the inclusion of the whole study of adaptationin "palaeobiology".It may happenthat a
r0)It is thereforenot homologousas a whole with the primary n.rantle,as shown earlier (Naef 1913, p.
180 388).The demal mantleis conservedas vestigesin
are generallyembeddedin sand,which explainsthe
different iocations,at leastin the Decapoda(cf. e.g.
betterpreservationof the rostrum.
Fig. 58), in particularin the nuchalarea.wherethe decapodanshell is still presentin the anteriorend of the rnantle(Fig. 10).
r2) Among the living Sepiinae,the generaSepiella, HentisepiusandMetasepiastand apart, as probable secondaryvariants of the Sepla type, from which
1t) The adhesiveconnectionin the nuchal area and the
they are not very distant.At any rate, they show no
similar funnel-mantle connectionsin Decapoda
closerelationshipto the older Belosepiinae,so we
functionas gliding surfaces(Figs 40 and 64).
can leavethem asidesincetheir link to the ancestral forms of the family must soughtthrough the living
1r) En.rbryosof octopods with well developedfins
genusSepla.
(Ciroteuthoidea)were not yet available. tt) He writes: "55 years ago Voltz has perfectly \ 3 )B e l o s e p i a t h u s c a n n o t b e c o n s i d e r e da f o r m
demonstratedthat the outer plate correspondsto the
intermediatewith the belemnites.as suggestedby
rostrum of belernnites,the internal part represents
Lang who oflered a rather arbitrary figure to
the phragmocone,in that the lamellae of the hump
illustratehis point of view (seebelow. p. 82).
continue posteriorly into the lamellae of the fork, the bent posterioredgesof the hump lamellaemust
]a) The authorsapparentlymissed the curvatureof the
be viewed as septal necks, and the reason why
initial part of the phragmocone.[Curvature]may
belemnitephragmoconesoften fall out of the rostra
still be recognizableevenin the distalpart.
is probably related to the epicuticula of the phragmoconesthat correspondsto the rnedian
15)Deshayesattributesthe speciesto Blainville whose publicationis dated1825(cf. p. 56).
plate". Riefstahl'stheory of shell growth by intusception haslong beenshownto be enoneous.
16)That is the phragmocone! ra)Here belong:S. aculeatad'Orb. 1839,Pl. 5 bis, Pl. 17)That is the periostracum!
25, Fig. 4; S. rouxii ibid. Pl. 19,Fig. 7; S. blainvillei i b i d . P l . 2 1 , F i g . 4 ; S . r o s t r a t ai b i d . P l . 2 6 e t c ; S .
1E)Probablynot quite as much (Fig. 23).
microtyledon Ortmann 1890 (very high fork!); S. .frameaOftmann 1890;S. koenlitziHoyle 1901.
1e)Errors of identificationcannotbe ruled out. Since acids in the environrnent(CO2?)seem to readily
r5)We now know that the genusTrachyteuthis("Sepia"
dissolve the rostrum and the sheathin sepioids,
hasti/orntis Riipp.) from Solnhofenhas nothing to
some fbssiliferous strata preserve only the
do with the sepiid family, striking similarities
phragmocones.It is therefore conceivable that
notwithstanding.
chamberedshellslike Spirulirostra can be mistaken for Spirtrla.Also seeunderSpinrliro.strina,p.T6.
2 6 ) S u b s e q u e n t l yr e n a m e d A r c h a e o t e u t h i s ( L e t h . Geogr.,vol. I, p. 520).
20)cf. Naef, Cephalopoda, vol. I, chapter39. ']7)This view was expressedby Prof. Rollier (Ztirich). 2t) cf. Zrttel, Grundzrige,editions of 1915 and 1921. Sacco(1904.p. 6) supposes this form to be merely
r8)SeeBlainville's(1827)opinion,p. 56 and 82.
basedon isolatedphragmocones of Spirulirostra. As in the case of the latter genus,the sheathsof
re) Teu0ro :calamary (Aristotle), calarnari is derived
Spirulirostrina are indeedoften destroyedby the
from Calamarium(It. Calamaio)writing tools (pen
acidity of the marl, so that only phragmoconesare
and ink). The pen (Calamus,Gladius) and the ink
preserved.However, the specimensof Spirulirostra
sacarehousedin an envelope,the mantle.
l8l The scopeof this group was essentialiyrecognized
an ancientLoligo which looks so similar to the pen
b y d ' O r b i g n y ( 1 8 3 9 ) w h o c a l l e dt h e m L o l i g i d a e "
of the living Loligo sagittata that I named the
before dismemberingthe group in an arbitrary u,ay
specimenLoligo sub.sagittato. To my knou'ledgeit
( 1 8 4 5 ) . K e f - e r s t e i n ( 18 6 6 ) a g a i n u n i t e d t h e
is the only specimenfrom the Solnhofenbedsthat
"DecapodaChondrophora"(p. 1441) and limited
can be secureiyplaced in the genusLoligo. The
the subdivisionsMyopsidaeand Oigopsidaeto this
other horny pens in shape reminiscent of a
group, but then mixed them up with true sepioids (Sepiolidae).
triangularsword (p. 583), which up to now have been generallyconsideredto be the internalhorny pens of Zollgo, probably belong either to the genus
'r0)An exceptionare someRecentOnychoteuthidaeand
Ottvchoteuthis or to yet anothergenus since they
Enoploteuthidae,in which all the radular teethhave
have small hooks ("griffes ou crochets")on their
only onecusp,as in the Sepioidea.
arms insteadof circular suckers.Of the 20 species olSepla-like cephalopodsin my collection.I have
3r) Sometimeshe did, at othertirneshe did not assume
ordered illustrations to be made of the most
that a phragmocone was present,cf. p. 169and 108. -'2)Milnster calledthern"fins".
noteworlhyonesso as to make them betterknor.vn. 3e)Eichstiitt,Solnhofen,Daiting,Nusplingen.
= \1 Onychoteuthi,s prisca Miinster 1828,basedon shells
'") Aentoo: slender.This is the largestfossilsquid.
from the Upper Lias "in the shalesnearAalen, Bo11,
=
Steiningen,Ohrndenand other places",which he
ar) The posterior part of this form was arbitrarily,
apparentlyidentifiedwith the prototeuthoidshells
indeedemoneouslycompletedon the basisof a very
= =
which he knervfrom Solnhof-en.
roughly sketched fragn.rent,which is obliquely
=
compressed. }) Of coursehe noticedthe inconsistency and tried to somehorvsolvethe problem.He thus offered(1836,
+ r ) V o l t z ( 1 8 3 6 . p . 3 2 4 ) s p e a k so f " g r o w t h l i n e s
p.325) a strikinglyvagueidea abouta dorsaland a
representingthe ends of a series ol mutually
ventral membranethat could have generatedthe
superimposedsheets".He also draws attentionto
median and lateralplates.The samecan be noted
tlre diff-erencefrom Loligo, in that accretionoccurs
for the deviation in the growth lines of the sheath
at the blunt end rather than at the pointed end. We
(in the alveolus)and on the outer surface.The pro-
see here the contrastbetween Prototeuthoideaand
ostracumis thoughtto inserl on the sheath.
Metateuthoidea.
-'5)Of coursethe threelayersare againdistinguishable!
ar) This pumpkin-sizeconcretionwas found in the Lias
16)ll),r1oroo : closely related.namely (seemingly)to
e l L o w e r T o a r c i a n ] n e a r S c h c i n e b e r gb y t h e ReverendHaftmann;it was he who split the nodule.
recentteuthoids(ommatostrephids).
= 't)
-") Mrinster 1846 (p. 57). Subgenus"Dor.,-anthes" for
Referring to the keel, Quenstedt (1849) characterizedthe Beloteuthidae- in particular- as
Acanthotetttil.swith arrow-shapedanteriorend of
"Crassicarinati"as opposedto the "Tenuicarinati"^
shell. SubgenusAcanthopusfor gladiuswith simply pointedend.(Incompletespecimens).
r'vhichcomprisedthe prototeuthoidsknown to l.rirn, excludingPlesioteuthis.The latterwas placedin a third group named"Hastiformes".The fourth group
r3)Mrinster(in letter to Bronn) reportsp. 582: "Some
containedhis genusSepia(Trachvteuthis).
time ago I found. in the natulal history collections o f t h e D u k e o f L e u c h t e n b e r ga r E i c h s t A t r ,t h e feather-shaped, horny gladius [Gcr. Leistchen]of
a5)The sameprobablyappliesLo Geoteuthi,s, Belopeltis. P arapIesioteuth is (?), andB eIoteuth i.sl
=
r82 ro)
Reuss: Loliginidenrestein der Kreideformation.
o f p h r a g m o c o n e so f 8 .
Abh. D. k. bdhm. Ges. d. Wiss. (5), vol. VIII,
i m p r e s s i o n so f t h e p r o - o s t r a c u m ,w h i c h w e r e
Appendix p. 28, plate,figures 1-2.
reproducedby Buckland(1836,Pl. 44') ( cf. Boud
semisulcatus with
1832). Their identihcation was incor:rect,however. ,r')
Fritsch, A.:
Cephalopoden der
bohm.
They belong to Acanthoteuthisspeciosa(Fig. 91),
Kreideformation 1872 (with the collaborationof U.
which in turn could be identical with B.
Schliinbach).
(cf. Fig. 90). (?) semisttlcattrs
a8) Quenstedt(1830, p. 163) had already noted that Zieten's variant of Loligo bollensisin Pl. 37, which
56)Agassiz (1835, Jahrb.,p. 168) writes: "My trip to E,ngland provided me with some important
is the earliest known Beloteuthis, representsa
information about the organisationof belemnites.I
separategenus.
have found that the so-calledOnychotettthisprisca with ink sac, as illustratedby v. Zieten (as Loligo,
ae)Regarding Schtibler see Alberti 1826: Uber die
Pl. XXV), is merely the anterior prolongation of a
schwtibischenFliizformationen der Gebirge des
belemnite,namely B. ovalis, as shown by an intact,
Wiirtternberg. Konigreiches Slultgart.
p e r f e c t l y p r e s e r v e ds p e c i m e na c c o m p a n y i n g3 5 new speciesof fishesfrom the Lias at Lyne-Regis,
50)lt has been collected in Germany, as well as lrom
seenin the collectionof Miss E. Philpot.Thus the
the SwabianLias, at Hondelagenear Braunschweig
belemnites have the anterior prolongation of the
and at Scheede(nearthe Mittellandkanal).
alveolus in the form of the plate of Onychotettthis and the ink sac of Sepiainside.The belemnitesthus
5r)Ke).o,Lvro - harpy,witch.
differ from Sepia marnly in showing a much stronger developmentof the point at the upper
52)So far, Pctlaeololigocould be included!
margin of the so-calledcuttlebone!lf the genera thus coalesce,what will happento the speciesonce
s3)SeeCephalopoda,vol. l, p. 122.
we know exactly how the different stagesin the growth of an individual come about?".
5a)Our knowledge is particularly fragmentary in this area.Gladii are well preservedonly in very specific marine sediments;significant inforrnation can only b e o b t a i n e d f r o m l i m e s t o n e sa n d m u d s t o n e s .
") H. v. Meyer (1836,p. 55) alsomentionsdrawingsof shells from the Lias of Lyme Regis, which Bucklandbroughtto a meetingin Bonn.
According to E. Stolley, there is a similar rock called "Tock" on the island of Helgoland;its
5E)These fossils are particularly useful as a direct
geologicalage is doubtful, however.Perhapsnew
confirmation of the general insight which was
data will be found in it. Apparently some shells
gained indirectly, following Voltz (p. 168), from
from this formation are housed in the Hamburg
the growth lines of belemnite phragmocones(cf.
museumof natural history. See Dawkins (1864) on
Figs 71,73 and 90). In particularthey allow us to
a questionablefragment of Beloteuthls from the
observedirectly the presenceof a pro-ostracumon
EnglishRhaetian.
the phragrnoconeand to determine its relative length.
55)This is a broken phragmocone,which containsa displaced ink sac; its end is furnished with a
5e) Unfortunately we only know the embryos of
paxillose belemnite rostrum (8. ovalis). The
polypodoids,in which extreme reduction of the
ostensible pro-ostracum is apparently only the
shell has taken place. This fact thus carrieslittle
conotheca.At that time, most peoplehad no precise
weight in our discussron.
knowledgeof the relative size of the phragmocone, a l t h o u g hM i i n s t e r ( 1 8 3 0 , P l . I , F i g . 1 5 ) , w i t h o u t fully understandingthe structures,gave illustrations
60) Miinster (1828) did not apparently distinguish teuthoidsfrom belemnoids.His "Onvchoteuthis"is
r83 said to occur in the Upper Jurassicof Solnhofenas well as in the Lias of Swabia(p. 579-581).He later
"o) See Figure 68! The hooks have a characteristic shape.I can neither confirm nor refute the presence
calledthe speciesfrom the Ltas "Acanthoteuthis",
of suckers.In rny view the impressionsare no1
but he also ur.ritedbelemnoidsundel the samename (4. speciosa : ./brrussacii - lichtensteinii) as
sufficientlydistinct(cf. p. 29).
teuthoids(Plesioteuthis)in which he erroneously
67)Zieten (1830) also studiedand figuredthe loliginid
assumed the presence ofhooks.cf. p. 181.
shell and its relation to the soft parts and described
Sternberg(1820) alreadyknew the arm crowns of
their animals;thus he indirectiyexplainedthe role
Ac. .speciosa.He describedthem as "Caulerpes
of the pro-ostracum.
the general connectionbetween fbssil gladii and n ' )
princeps", mistaking them for plant remains(green a l g a e ) . M i i n s t e r ( 1 8 3 4 ) i n t e r p r e t e dt h e m m o r e correctly basedon his knowledge of On.vc'hoteuthis Lichtenstein.Onp. 42 he reportson two speciesof "cuttlefish", one of which had arms with small
6x)Not even passivelyunder water pressure. good In swimmersthey areprobablyrelativelysmaller. i'e)He views it as a weapon,much like clarvs!(?)
suckers.The latter are said to be S-shaped:"it thus appearsthat these cephalopodsfrom the Jurassrc
'-0)Precisecalculationsare not in generalpossible (cfl
differ from living cephalopodsby the shapeof their
Abel 19i6, p. 166);they rvouldbe rrost desirable.
suckersas much as the frshesdiffer from later ones
They would have to be based on reliable
by the shapeoftheir scales".(The "fins" and "taillike process" are interpretedas artefacts of
information on the relative sizes of the phragmoconeand rostrum,thicknessof the sheath
fossilisationof the mantlesacand phragmocone).
and conotheca, massof the siphuncleand the septa. Taking everything into account, the buoyancy
ot)
Up to now the general assumptionwas that there were 8 arms. and Ac. spec'iosathereforewas often placed in the Octopoda, especially by Miinster ( 1 8 3 7 "1 8 4 3 ) ,R . W a g n e r( 1 8 3 9 ) ,B r o n n ( 1 8 4 8 ) , Rcimer(1852),and so on up to Btilow (1920).
should be somervhatlower than that supposedby Abel. but still too great to permit permanentliie in deepu.ater. rr) If indeeda Celaeno-Itkelife form must be assumed for Acanthoteuthi,s problematica(p. l8a), it could
u : ) The hooks do not all show the same degree of
only be that of an atypical. benthic variant of the
curvature;somelook more like claws usedfor mere
belemnoids.The exceptionwould then confirm the
scratching,but the tenninal parts may have been
rule, in demonstrating how lar the rnoditicationof
lost. Figure 63g showsa very completespecimen,
the type must proceedto achieveadaptationto life
which apparentlywas able to seizeobjects.At the
on the bottom.
arm basesthe hooks are rather small, they then progressivelyincreasein size,and distallydecrease againand finally disappearfrom the picture.
rr) At best the stocky genusSepioleuthi.s. which I have not seenalive, may be an exception.The sepioids have apparentlybecome adaptedto benthic life
nr)
In Celaenothe shell itself is well preserved,u,hereas
through a ,seconclarl,modification of the shel1
in Acanthoteuthis Ihe conothecaand pro-ostracum
apparatus.
are completelymaccratedand ahnost completely dissolved. t,-,)
In Celaeno conic'u the grovr,thlines are closely
13)It is not correctto use the term "alveolus" for the phragmoconewhich fills that space;early authors
spaced,whereas in the present fbrm the u'idely
sometimescalledit the "'alveolite".Here I can only give a very generalview of the variablecollcepts
spacedparallellines on the phraemocone probably
and terms of belernnoidmorphology.One of the
representsutures.Parts of thc phragmoconeare
aims of our u.ork is to provide more precisionand to establishthe meaningol technicalterms in our
shatteredand thus cannotbe reasscntblecl.
=
t84 8r)Lias (!
language. ra) This apparentlyapplies also for the dorsal grooves, S e eL i s s a j o u s( 1 9 1 5 ,P 1 . ".). 1, Fig.2 showinga clearly similar slit field in the
Ea)In Swabia!
a t i e a s t n D i c o e l u s( q .
E 5 )F o r t h e J u r a s s i cb e l e m n i t e ss u c h a n a n a l y s i s
dorsaland the ventralpart,respectively).
apparentlywas carriedout by the late M. Lissajous; the resultswill be publishedby the University of
15)The embryos of cephalopodslive in a delicate
Lyon. For the Cretaceousbelemnites it can be
globular chorion. To perforatethis envelope,some
expectedfrom ongoingwork by E. Stolley.
forms have specialterminal spines(p. 98). Mature hatchiings always leave the egg caseposterior end
E 6 )S t o l l e y ( 1 9 1 9 , p . 3 5 ) s u p p o s e st h e e x i s t e n c eo f
first. cf. Naef, Cephalopoda,vol. II, chapterson
intermediateforms and draws attention to the rich
Sepiolidaeand Octopoda.
belemnite material from the mines near Harzburgin the foothills of the Schlewecke-Harlingerode
76)lts special characteris probably reflected by its
Harz mountains.The connectionhe has in mind
capability to become secondarilydissolved,so that
would involve the oldest Paxillosi (p. 234) via
a longitudinalcanalis formed (8. perJbratus).
thick-stalked "Clavati",
especially B.
charmouthensis. I wonder whether they belong 7') In shells of recent cuttlefish it is easy to neatly
here. Unfortunately the juvenile rostrurn is
separate the shell parts corresponding to the
unknown.
phragmocone(usinghydrochloricacid in alcohol).
\-) Stolley ( 1919) united these forms with the 7s)Note the striking fact that Abel (1916) doesnot even
Hastitinae and Coeloteuthinae.We exclude the
attemptto reconstnrctan entire belemniteshell and
latter subfarnilies;according to current rules of
to clarify its morphologicalrelationships.
nomenclature(p. l9) the namesof subfamiliesand families shouldbe basedon the typical genera.
re) Here is a potential spherefor an ecology of extinct organisms,which would not make sense as an
*n)
independentdiscipline. Dollo, the lounding father
Quenstedt(1830, p. 166) regardedthis speciesas young rostra or apical parls of old specimensof ".8.
of such a tendencywithin modern palaeontology
giganteus".
consistentlyusesthe older tenn "ethology",whose general use has been pushed aside by Haeckel
*o)
The similarity is also striking when comparing
( 1866). On the other hand. Abel's tern-r
globularvarietiesof rhenanato S. sulcata or S. raui
(Palaeobiology)is ratherunfortunateat a time when
( c f . W e r n e r1 9 1 2 , P l .1 2 ,F i g . 2 w i t h P l . 1 1 ,F i g s 7
Biology meansthe generalscienceof life, at leastin
a n d 8 ) . T h e r e l a t i o n s h i pw i t h O d o n t o b e l u s t s
standardliterature.
doubtlesscorlmon.
E0)Abel's (1916,p. 188)idea aboutanirnals"ploughing
e 0 )C o m p a r em y F i g u r e s6 2 , 6 3 x , y , 6 7 b , 9 0 a n d 9 1 ,
through the Posidonid grass weeds" is out of the
w i t h Z i t t e l 1 8 8 5 ,p . 5 1 1 ,F i g . 7 1 3 ;p . 5 2 0 , F t g .7 1 4 ;
question!
ibid. Crundziige 1921, p. 587. The pro-ostracum (from Solnhofen) figured by Zittel is the same as
8r) Such (unfotunately frequent)conclusionsare due to Haeckel's so-called "fundamental law biogenetics". SeeNaef (1917,p. 61;'1920).
the one hereshown(in greaterdetail)in Figure 90.
of er)I have not been able to find this specirnen;I lear it is a misidentified shell of Acanthoteulftlswithout a
82)Lias yl
rostrum. cf. figures in Mtinster (1830), Buckland (1836),Quenstedt (1849).
185 ot)
ln Aulacocera,sthe siphuncle deviates from its
vh)Perhapsthereis one exception:d'Orbigny(1839,Pl.
marginalpositiontou'ardsthe posteriorend (in the
15. Fig. 6) describedand figured a "Sepia " venusta
secondchamber),to enter the protoconchin the
Mtinst. (1837, p. 252) from Upper Jurassicsrrata
centreof the first septum(,',.Biilow. 1915.p. 33).
(seealso d'Orbigny.1846,Pl. 5. Fig. 7. and Chenu,
This seems to be the normal situation in
1 8 5 9 .p . 4 5 , F i g . 1 3 7 ) .T h e s p e c i m e nc a n n o tb e a
d i b r a n c h i a t e s ;w e f i n d i t i n t h e o n l y l i v i n g
Sepia, any more than a Trac.hyteuthis, which is
representativeil,hich has conservedthese parts
classifiedwith Sepia by d'Orbigny. In contrast,the small shell (preservedin Munich) shows some
v i r t u a l l y u n c h a n g e d( c f . S p i r u l a p . 4 1 , F i g . 9 ) . I n neithercasedoesthe siphunclereally communicate
sirnilarity to the juvenile brood shell of Argonautcr (cf. Hoyle, 1904).
rvith the flrst char.r.rber. The initial caecum rn Spirula is covered only by a weakly calcified c o n c h i o l i n c a p w h i c h i s n o t r e c o g n i z a b l ei n
er) This tendencyis systematicallyexploited by the
belemnites.
fishermene.g. of southentItaly; they string together a seriesof pots and lower them to the sea bottom.
,,) Hauer's repeatedassertionthat the siphuncle of
When they are brought up after some time, most
Atrlacocerasis dorsal(cf. Pompecky1912,p. 296c)
pots are occupiedby octopusesclinging to their
did not convinceme and inducedme (1912,p. 250,
hiding-p1aces.
Fig. a6) to assumea central position of the siphunclefor the ancestralforms ol the belemnites
er) Joubin (Bull. Monaco.No. 351, p. 2) c r e a t e sa
or the belemnoidsand the dibranchiates in general.
separate.certainlyuntenablefarnily. for this forrn
This turned out a gross error. but at the time of its publicationit u'asundulycriticised.
u.hi ch appalentl,vresentbles the enoploter-rthids. '/")
,t)
Erroneouslyidentifiedas "B. ov'enil Pratt" (p. 65)
Here belon,ss"C'hirotetttltis"portieri Joubin 1916 ( B u l i . M o n a c o .N o . 3 1 7 ) . T h i s i s a v e r y p e c u i i a r
and confusedwrth Belemnitespuzosiarzr.s d'Orb.
oegopsid, which has not rnuch to do u'ith
frornthe samelayers(Zittel 1885,p. 501,Fig.68a).
Chiroteuthis; indeed it cannotbe placed close to any known genus.The overall aspectremindsone
,rS )
This may be due to the delicate structureand small
of larvae of Abroliopsis. Seeoriginal descriptionby
size of the ancestralforms, which I envisageas
Joubinand Cephalopoda, vol. I (chapter9,1923).
swirnming and crawlir.rganimals: such a drastic degenerationof the shell is imaginableonly in a seriesol lbrms in u'hich shell developmentu,'ent barely beyond the first shell rudirnent. Thus it
ttt1lBolitaenadiaphana(Chun 1915,p. a93) should be consideredas the nominal type of this group of genera;the distinctive feature is the rnulticuspid
also seemsnatural that the earliestsepioidsu'ere
form of the lateral teeth of the radula. which erist
small forms, wirose relatively sirnple structure p e r r ni t t ed f a r - r ea c hi n g s h if t s i n p r i m a r y
aisoin Amphitretus (loc. cit., p.533). The
relationshipslof parts]. Conversely,increaseof body size seems to accompanyelaborationand
to any form of radula, so that it is still valid when
improverlentol establishedtypesof organisation.
designationis sufficientlygeneralto be applicable closerelativeshave a somewhatsimplerform.
187 (s04)
d u s u d - o u e sd t e l a F r a n c e .A c t e s S o c . L i n n .
Referencelist
Bordeaux,t. 42. 1895. Bemard,F., Elernents de Paldontologie. Pans.
1916. Abel. O., Paliiobiologie der Cephalopoden. Jena, G. Fischer.
1900. Biedermann,W.. Untersuchungenriber Bau und E , n t s t e h u n gd e r M o l l u s k e n s c h a l e n . J e n .
1921.Ibid., Die Methoden der paliiobioiogischen F o r s c h u n g .H a n d b . d . b i o l . A r b e i t s m e t h o d e n (Abderhalden).
Zeitschr.,Vol. XXXVI, N.F. XXX. 1861.Binkhorst, J. T. van den, Monographie... c o u c h e sc r e t a c 6 e ss u p d r i e u r e sd u L i m b o u r g .
1835. Agassiz,A., Uber Belemniten.Leonh.& Bronns Jahrb.ftir Min.
C d p h a l o p o d e sp . 1 - 1 2 . 2 . e d i t i o n . B r t i s s e l MaestrichtI 873.
1905. Alessandri,G. de. Avanzi di un nuovo generedi Cefalopododel Eocenedei dintorni di Parioi
1 8 2 5 . B l a i n v i l l e , D . d e . M a n u e l d e M a l a c o l o g i ee t Conchyliologie.Paris 1825-27.
Riv. Ital. Paleontologia, v. XI.
1 8 2 7 . I b i d . . M d m o i r e s u r l e s B e l e m n i t e sP . a r i s .( c f .
1902. Angermann.E,.,Uber das Genuslc'clrlhoteuthis Miinst. aus den lith. Schiefern Bayerns. N. Jahrb.f'. Min., Suppl.Vol.. XV. 1887. Appelidf. A., Om skalets bildning hos Sepia o/ficinalisL. Ofvers.Vet. Akak. Forh. 1894. Ibid., Die Schalenvon Sepia, Spirula und N a u t i l u s . K o n g 1 . S v e n s k aV e t . H a n d i . . V o l . XXV.
Nouv. Bull. Soc.philom. 1825and Ann. Sc. nat. t. vrl. 1826). 1882. Blake, Monographof British Cephalopoda. Part l. London. 1 8 6 1 . B l a n f o r d , H . . T h e f b s s i l C e p h a l o p o d ao f t h e cretaceons rocks of SouthernIndia. Mem. Geol. Surv.lndia. Calcutta. 1844. Bou'erbank.J.S.,Observationson the Structure
1899. Ibid., Uber das VorkommeninnererSchalenbei gen Cephalopoden(Oc'top oda). Bergens achtarr.ni Mus. Aarb.No. 12. 1911. Arthaber,G. v.. Die Trias von Albanien. Beitr.
olShells.Trans.Micr. Soc.London. 1880. Branco.W.. Beobachtungen an Aulacoc:erasy. Hauer.Zeitschr.d. D. geol.Ges.,Vol. XXXll. 1879/80.Ibid.. Beitriige zur Entu,icklungsgeschichte
z . G e o l . u . P a l . O s t . - U n g .r r . d . O l i e n t s ,V o l .
der lbssilen Cephalopoden. Palaeontographica,
XXIV.
Vol. XXVI u. XXVII.
1857.Barrande, Uber die innere Struktur der Nautilidenschalen. N. Jahrb.f. Mikr. 1870. Ibid., Distribution des Cdphalopodesdans les contrdessiluriennes.Extr. du. Syst.sil. du centre de la Boh6me,v. II, Texte V. Pragueet Pans. 1909. Bauer, V.. Einfrihrung in die Physiologieder Cephalopoden.Mitt. zool. Stat. Neapel. Vol. XIX.
f o s s i l e n C e p h a l o p o d e nZ. e i t s c h r .d . D . g e o l . Ges. 1 7 3 2 .B r e y n i u s , J . P h . , D i s s e r t a t i o p h y s i c a d e p o l y t h a l a m i s , n o v a t e s t a c e o r u mc l a s s e Gedanie.(Danzig.) 1 8 8 0 .B r o c k , J . , V e r s u c h e i n e r P h y l o g e n i e d e r dibranchiatenCephalopoden.Morph. .Tahrb..
1878. Bayle, E., Expiicationde la cartegdologiquede l a F r a n c e .t . 4 ,
1880. lbid., Uber die Verwandtschaftsverhiiltnisse der
Atlas lre partie: Fossiles
p r i n c i p a ud x e st e r r a i n s . 1838. Bellardi, L., Notice sur I'Argonauta nitida Lk. B u l l . S o c .g e o l .F r . ( l ) t . g . 1872.Ibid., J. Molluschi dei terreni terziari de1 Piemontee della Ligr-rria.p. l. 1910. Benecke,E. W., Uber Belr'nrttiteslurc.sulac'tus. Zentralbl.f. Min. 1 8 3 1 . B e n e t t .G . , T h e i n h a b i t a n to f p e a r l y N a u t i l t r s . London medic.Gazette.v. VIII. 1888. Benoist.E. A., Descriptiondes Ciphalopodes... Coquillesfossilesdes terrainstertiairesmoyens
Vol. VI. Broili, F., cf. Zittel Grundzrtge1885. 1858. Bronn. H.G., Beitriigezur triasischenFaunader bituminosenSchieferi. Raibl. N. Jahrb.f. Min. ( c f . i b i d .1 8 5 9 . ) 1852. lbid. & Romer,F., Lethaeageognostica, Vol. Ill. 2nd edition Stuttgart. 1 7 8 9 . B r u g h i e r e "E n c y c l o p 6 d i em d t h o d i q u e .P a r i s . Fortsetzr"rng: DeshayesI 830. 1829.Buckland. W., On the discoveryof a new s p e c i e s o f P t e r o d a c t y l ea n d . . . o f a b l a c k substance r e s e m b h n gS e p i a . . . . i n t h e L i a s o f Lyme Regis.Geol.Soc.London.
188 1 8 3 5 . I b i d . . N o t i z i i b e r d i e h y d r a u l i s c hW e irkungdes Siphosbei denNautilen...N. Jahrb.f. Min. 1836.Ibid..
Bernerkungen iiber
das
Genus
Belentnoseplaund den fossilen Tintensackin demvorderenKegelderBelernniten.Ebenda. 1836. Ibid., Geology an Mineralogy consideredwith referenceto natural Theology. (,.Bridgewater Treatise")London. (c1-.Agassiz.) 1 9 1 5 . B r . i l o w ,E . v . , O r t h o c e r e nu n d B e l e m n i t e nd e r Trias von Timor. Paliiontologiev. Timor, 4. Lief. Stuttgafi.
from
the
Upper
Lias
Alderton,
of
Proc.Mal. Soc.,v. II (cL v. I). Gloucestershire. 1 8 9 7 . I b i d . , A c a n t h o t e u t h i s, s p e c ' i o , sMai i n s t . G e o l . Mag. (4), v. IV. 1897.Ibid., On an example ol Acanthoteuthis speciosa.Ibtd. 1900. Ibid., The buccal membraneof Acanthoteuthis Proc.Mal. Soc.,v. III. specio.sa. 1 9 0 2 . N o t e o n t h e T y p e - S p e c i m eonf B e l e m n o t e u t h i . s Montefiorei.J. Buckmann...Proc.Mal. Soc..v. V.
1916. lbid., Uber einenPhragmoconvon Aulacoceras strlcaltmtvon Hauer aus der alp. Trias. Zentralbl. Min.Geol.Pal.
1902 Ibid.. Note on a Dibranchiate Cephalopod (BelopterinaLevesqueid'Orb). Ibid. 1 9 0 4 . I b i d . . C e p h a l o p o d sf r o m t h e N . W . I n d i a n
1920. lbid., Fossilium Catalogus.Animalia. Pars 11. Berlin.W. Junk. Cephalopoda dibranchiata. 1894. Bullen, N.R. & Harris, S.F.,A Revisionof the British Eocene Cephalopoda.Proc. Mal. Soc. London,v. 1.
Frontier.Geol.Mag. (5), v. L 1907. Ibid., On the ams of the Belemnite.Proc. Mal. Soc.v. VII. London. 1910. Ibid., On BelemnocamaxBov'eri n. gen. et sp. lower chalk oflincolnshire. Proc.Geol. Ass.,v.
1892. Canavari,M., Note die Malacologiafossile.2. S p i r t t l i r o s t r i nLao v i s a t o i n . g . , n . s p . d i e
XXI. 1 9 1 5 .I b i d . ,
A
Dibranchiate
Cephalopod
Cetalopodoraccolto nel terziario Sardegna...
(Plesioteuthis)from the Lithographic Stone
Bul. Soc.Ma1.Ital.,v. XVI. 'W., 1843. Carpenter, On the microsc. Structure ol
(Lower Kimmeridgean)ol Eichstiitt,Bavaria. Proc.Mal. Soc.,v. XI. London.
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1 9 1 8 . I b i d . , R e c e n t r e s e a r c h e so n t h e B e l e m n i t e
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1 8 9 8 .G r i f f i n . E . L . , T h e a n a t o m y o f N a u t i l u s Pompilius.Men. Nat. Acad.Sc.,v. VIIL 1 8 6 1 . G i i m b e l ,C . W . v . . G e o g n o s t i s c hBee s c h r e i b u n g desbayerischenAlpengebirges. 1 9 1 5 . H a n i e l .C . A . . D i e C e p h a l o p o d edne r D y a s v o n Timor. Wanner.Pal. v. Timor. Stuttgafi. 1860.Hauer" F.v., Nachtriige zur Kenntnis der Cephalopodenlauna der HallstiidterSchichten. S i t z . - B e r .A k a d . W i s s . W i e n , m a t h . - n a t .K l . , Vol. XLI. 1892. lbid., Beitriigezur Kenntnis der Cephalopoden aus der Trias von Bosnien:I. Neue Funde aus dem Muschelkalkvon Han Buloy bei Serajevo. Denkschr.Kais.Akad. Wiss.Wien, Vo1.LIX. 1865. Hebert,F., Sur le groupde Belentnites auquelde Blainv. et d'Orbigny ont donn6 le nom de B. breuls.Buil. Soc.Gdol.France(2), t. 22. 1 8 7 7 .H e c t o r . O n t h e B e l e m n i t e s f o u n d i n N e w Zealand.Trans.Proc.New. Zeal. Inst.,v. X. 1 9 1 5 . H i l b e r , V . , D i e e i l t e s t eb e k a n n t e u n d e r s t e miociine Argonauta. Mitt. naturw. Ver. f. Steiermark,Vol. Ll. Graz. 1903. Hoernes,R., Zur Ontogenieund Phylogenieder
190 C e p h a l o p o d e nJ.a h r b .k . k . G e o l . R e i c h a n s t . , Vol. Lil. cf. Biol. Zentralbl.,Vol. XXIIL 1886. Hoyle, W.E., Cephalopoda. ChallengerRep., v.
1882. Ibid., Dte Gastropoda ..., Cephalopoda und Pteropoda des norddeutschenMiociin. (II. Teil von: Das nordd.Miociin u. s. Moll.-Fauna.) ;a
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1 8 8 5 .I b i d . , U b e r d i e p a l i i o c i i n e F a u n a v o n
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seineMollusken-Fauna.IV. Berlin. 1825. Krinig,Iconesfossiliumsectilis.t. 2. 1801.Lamarck, J.B., Systdmedes animaux sans vertdbres.Paris. 1 9 0 0 .L a n g , A . , L e h r b u c h d e r v e r g l e i c h e n d e n
1872. Hyatt, A., Fossil Cephalopoda.Embryology.
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2nd edition,revisedby. K. Hescheler. 1907. Langerhahn,A., Mitteilung an Henn Joh. Bohm. Zeitschr.d. D. geol.Ges.,Vol. LIX (1906).
naturf.Freunde,Berlin. 1899. Ibid. (Kriechspurvon Acanthoteuthis).Z. d. D. geol.Ges.,Vol. LI.
1906.Leriche, M., Note sur le Genre Vasseurro Munier-Chalmas.Bull. Soc. Sc. Nat. de I'ouest
1902. Jaeckel,O.. Theseniiber die Organisationund L e b e n s w e i s ea u s g e s t o r b e n eC r ephalopoden. Ibid. Vol. LIV.
de la France(.2)t. 6. Nantes. 1 9 1 2 . L i s s a j o u sM , ., JurassiquM e a c o n n a i sF . ossiles caractdristiques. Bull. Soc. Hist. N. Mdcon. v.
1904. Ibid., Neue Beobachtungen an Orthoceren.Ibid. Vol. LV. 1891. Janet,Ch., Note sur trois nouvellesBelemnites sdnoniennes. Bul. Soc.96ol.Fr. (3), t. 19. 1896. Jatta, G., I Cefalopodi viventi nel Golfo di N a p o l i ( S i s t e m a t i c a ) .F a u n a u n d F l o r a d e s Golfesvon Neapel.23. Monogr. 1892. Joubin, L., Recherchessur la coloration du tdgumentchez les Cdphalopodes.Arch. Z. Exp., t. 10.
III. 1915. Ibid., Quelquesremarquessur les Bdlemnites jurassiques.Ibid. v. lV. 1896. Lonnberg,E., Notes on Spirtrla reticulata Owen and its phylogeny. Zool. Stud. Festskrift W. Lilljeborg.Upsala. 1916. Loescher,V., Zum Bett desActinocamaxplenus Blv. Zeitschr.d. D. geol.Ges.,Vol. LXVil. 1898. Logan, W., The invertebratesof the Benton, Niobara and Fort Pierre groups. Univ. geol.
1 8 6 6 .K e f e r s t e i n ,W . , M a l a c o z o a c e p h a l o p h o r a . Bronns Kl. u. Ordn. d. Tierreichs,Vol. III 2. Heidelberg1862-66.
Surv. Kansas, v. IV. PalaeontologyPart. 8. Topeka. 1 8 4 8 .M a n t e l l , G . A . ,
1 9 1 0 - 1 3 .K i l i a n , W . , L e t h a e a g e o g n o s t i c a2. . D a s Mesozoikum,Vol. III, Kreide. Stuttgarl. 1 9 1 5 .I b i d . , C o n t r i b u t i o n d I ' d t u d e d e s F a u n e s
Observations on
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the
Oxford-clay near
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paleocrdtacdes de sud-ouestde la France.Mem.
Supplementaryobservations1850). (cf.: Ann.
pour servir d l'explic. de la carte gdol de la
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1851.Ibid., pertrifications and their teachings.
1170-75.Knorr, G.W., Verlustigungder Oogen ... 6 Teile in 3 Biinden, 190 kolor. Tafeln. 1 8 6 7 .K o e n e n , A . v . , B e t r a g z v r K e n n t n i s d e r norddeutschen
Tertiiirgebirges.Palaeontographica, Vol. XVI. (cf. 1865,Zeitschr.d. geol. Ges.,Vol. XVII, p. 42e.)
1863. Mayer-Emayr,K., Liste par ordre systdmatique de Bdlemnitesdes terrainsjurassiques.Journ.de
Amsterdam. Mollusken-Fauna des
London.
Conch.t. 1l. 1 8 6 4 - 6 8 .I b i d . , D i a g n o s e sd e B 6 l e m n i t e sn o u v e l l e s . I b i d .t . l 2 - 1 4 . 1883.Ibid., Grundziige der Classification der Belemniten.Zeitschr.d. D. geol. Ges.
191 1 8 8 4 .I b i d . .
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Vierleljahrsschr.nat. Ges.Ziirich. 1860.Meek, F.B. & Hayden tjber Phyttoteuthis, N a m e ! ) P r o c . A c a d . N a t . S c . P h i l a d e l p h i av, . XII.
des C6phalopodeset
sur les rapports
zoologiquesdes Ammonites avec les Seiches.C. R. paris. 1 8 8 0 . I b i d . , S u r l e g e n r eV a s s e u r i aB. u l l . S o c .g e o l . France(3), t. g. paris.
1 8 6 4 . M e e k , F . B . , S r n i t h s o n i a nC h e k L i s t . N . A m . C r e t .F o s s i l sv, . X X V I .
1 8 8 7 . I b i d . , D i v e r s eB e i t r i i g ei n F i s c h e r ,M a n u e l . . . (s.d.).
1876. Ibid.. A Report on the InvertebrateCretaceous
1828. Miinster. S., Graf zu, Versteinerungenvon
and Tertiary Fossiis of the r-rpperMissor-rri
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1832. Meyer, H.v., Palaeologica.zur Geschichteder Erde und ihrer Geschdpfe.Frankfur-ta.M.
1830. Ibid., Bemerkungenzur nriherenKenntnis der
1 8 3 4 .I b i d . . U b e r L e p t o t e u t h i s
n. G. Mus.
1 8 3 0 . I b i d . .B e m e r k u n g erni b e rd a s V o r k o m m e nv o n pteroclactl,lrl.r, voll fossiler Sepia und von
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Coprolithen in Deutschland.N. Jahrb.f. Min..
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1846. Ibid. (rJberTrochvtetrrhis). N. Jahrb.f. Min.
1832. Ibid., Beitriigezur petrefaktenkunde. l. Heft (in
1856. Ibid., Trachvteuthisensifbrmis aus d. lithogr.
2. editon 1843).3. Heft: 1840,5. Heft: 1g42,6.
Schieferin Bayem. Palaeontographica, Vol. IV. i886. Meyer,O., andAldrich, T.H.. The tertiaryFauna
Heft: 1843"7. Heft: 1846. Bayreuth.
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1832. Ibid., Nouvellesobservations sur les B6lemnites. B o u dM d m . G d o l .p a l . ( c f . 1 8 5 0p. . 2 9 5 . ) 1912. Murray.J. and Hjort, J., The depthof the Ocean London.
1 8 2 6 . M i l l e r , J . S . , O b s e r v a t i o n so n B e l e m n i t e s . G e l e s e na m 4 . A p r i l 1 8 2 3 . T r a n s . G e o l . S o c . London(2) v. I.
1 9 1 2 . N a e f , A . , C e p h a l o p o d a . H a n d w c i r t e r b .d . N a t u r w . . v o l .l l . J e n a . 1912. Ibid., Teuthologische Norizen Nr. I
1802. Montfort. D. de. Histoire nat. des Mollusques. P a r i s1 8 0 2 . 1808.Ibid.'
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Anz., vol. xxxx-xl-. 1 9 1 3 . I b i d . , S t u d i e nz u r g e n e r e l l e nM o r p h o l o g i ed e r
Conchyliologie
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classification methodique descoquilles.paris. 1854. Morris, J., A catalogueof British Fossils.(2. edit.) 1 8 8 2 . M o j s i s o w i c s , E . v . . D i e C e p h a l o p o d e nd e r
M o l l u s k e n I. I T e i l . E r g e b .u . F o r t s c h rd. . Z o o L Vol. III. 1916. Ibid., Systematische Ubersichtder mediterranen publ. Staz.zool.Napoli. Cephalopoden. 1 9 1 7 . I b i d . . D i e i n d i v i d u e l l eE n t r . v i c k l u nogr g a n i s c h e r
mediterranenTriasprovinz. Abh. k. k. geol.
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Reichsanst. Wien, Vol. X.
Jena.G. Fischer.
1 8 8 5 .I b i d . ,
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das
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Atrlacoceras..lahrb.d. k. k. geol. Reichsanst.. Vol. XXl. (cf. 1871). 1 9 0 2 ' I b i d . .D a sG e b i r g e u m H a l l s t a dA t .b h . k . k . g e o l . Reichsanst.^ Vol. VI, Suppl. 3. Daselbstauch: D i e C e p h a l o p o d e ni n d e r H a l l s t e i d t eK r alke. Suppl.l. 1 8 7 2 . M u n i e r - C h a h n a sE.. , S u r l e s n o u v e a u xG e n r e s Belopterina et BavanoteuthisB:Jtl.Soc. Gdol. ( 2 ) ,v . X X I X . P a r i s( 1 8 7 1 ? ) France t873. lbid., Sur le ddveloppement dr-rphragmostracum
1 9 1 8 .I b i d . . B e i t r r i g e z n r K e n n t n i s d e r f o s s i l e n Cephalopoden. Verh. Schrveiz.naturf. Ges..99. Vers.Ztirich. 1 9 1 9 .I b i d . . I d e a l i s t i s c h e M o r p h o l o g i e phirlogenetik. Jena.G. Fischer.
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192 derselben.Mitt. zool. Stat.Neapel,Vol. XXII. 1 9 2 1 . I b i d . , D i e C e p h a l o p o d e n .3 5 . M o n o g r . d e r ,,Faunaund Flora des Golfes von Neapel". L Lieferung,Berlin. Friedliinderund Sohn.
1846. Ibid., Paleontologieuniverselledes coquilleset des rnollusques.Paris. Zum Teil identischmit Moll. v. et foss., unter Weglassungder rezenten Formen.Fragment.Ebensoist die Paldontologie
1921. Ibid., Uber die DeutungbelemnoiderFossilien
dtrangdrenur als eine teilweise Wiederholung
auf Grund des Baues und der Entwicklung
der Pal. univ. mit vermindertenTafeln gedacht.
rezenter Tintenfische. Verh. Schweiz. naturf.
Beide Fragmente sind oft verquickt und
Ges.,102.Vers.Schaffhausen.
verwechselt.
1 9 2 1 .I b i d . , U b e r B a u u n d L e b e n s w e i s e d e r
1850-52. Ibid.,
Prodrome de
Paldontologie
tetrabranchiaten Cephalopoden.Vierteljahrsschr.
stratigraphique universeile des animaux
Zrirch.naturf.Ges.
Mollusqueset Rayonnes.Paris(v. I-II, 1850,v.
1922.lbid.,
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B e l e m n i t e n r o s t r u m s .E c l o g a e G e o l o g i c a e Helveticae,Vol. XVI.
1836. Owen,R., Cephalopoda. Todd's Cyclopaedia, v. I. London.
1877. Nathusius-Krinigsborn,Untersuchungentiber nicht zelluliireOrganismen.Berlin. 1 8 7 9 . N i c h o l s o n , H . A & L y d e k k e r , R . , 1 8 8 9 ' 1 ,A Manual of
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1843. Ibid., On Cephalopodswith Chamberedshells. Beeing the twentythird of the Hunterian Lectures delivered at the Royal Coliege of Surgeons. 1 8 4 4 . I b i d . , A d e s c r i p t i o no f C e r t a i n B e l e m n i t e s ,
1885. Niemiec,J., Recherches morphologiques sur les
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I 8 4 1. )
1890. Norman, A.m., Revision of British Mollusca. Ann. Mag. N H. (6).,v. V.
1 8 5 5 . I b i d . , N o t i c e o f a n e w s p e c i e so f a n e x t i n c t genus
1863. Ooster, W.A., Pdtrificationsremarquablesdes
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Alpes suisses.Cataloguedes Cdphalopodes des
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1862. Oppel,A., Uberjurass.Cephalopoden. Pal. Mitt. aus d. Mus. Kgl. Bayr. Staates.Stuttgart.(cf. Pal. Mitt. 1863and Wiirttemb.Jahreshefte 1855,
1899. Parona,C.F., Note sui Cefalopoditerziarii del Piemonte.Palaeontogr. Ital.,v. lV. 1842. Pearce,Ch. (Uber Belentnoteutlzls). Proc. Geol.
12.Jahrg.) 1839. Orbigny,A.d'(cf. F6russac1835). 1841 Ibid., Considdrations pal6ontologiqueset geologiques sur
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la
distribution des
Cdphalopodes acdtabulifdres. Ann. Sc. nat. (2), t. 16.Zoologie.
Soc.London,v. III. p. 593. 1 9 0 6 . P e l s e n e e r ,P . , M o l l u s c a i n : A T r e a t i s e o n Zoology, editedby E. Lankester,v. V. London. 1912. Pfeffer, G., De Cephalopodender PlanktonExpedition. Zugleich eine monographische
1842. Ibid., M6moires sur deux genresnouveauxde
Ubersicht der oegopsidenCephalopoden.Erg.
C e p h a l o p o d e sf o s s i l e s ( l e s C o n o t e t r t h i se t
Plankt.-Exp. d. Humboldt-Stiftung. Kiel u.
Spirtrliro.stra)offrant des passages,d'un c6td
Leipzig.
entre la Spirule et la Seiche,de l'autre entre les
1865-70. Phillips, J., A Monograph of British
Belemniteset les Ommastrdphes. Ann. sc. nat. ( 2 ) t . 1 7 .Z o o l o g i e .
1899. Picard.,K., Uber Cephalopoden ausdem unteren
1840-49.Ibid., Paldontologiefrancaise.Paris. (Terres jurassiques,I. 1., 1842- Temescretacees. t. 1., 1840 1842.) 1 8 4 5 . I b i d . , M o l l u s q u e sv i v a n t e se t f o s s i l e s. . . t . L Cephalopodesacdtabulifdres.Paris.New edition 1855 (nearlyunchanged).
Belemnitidae.Palaeontographic Soc.London. Muschelkalkvon Sondershausen. Monatsbl. d. D. geol.Ges. 1910. Ibid., Campvlosepiaelongatan. sp. Ibid. 1858-64.Pictet et Campiche,Materiaux pour la Paldontologie Suisse.v. I et II. 1902. Poeta, Ph., Uber die Anfangskammern der
193 Gattung Orthoceras Breyn. Sitz.-Ber. k. bcihm. Ges.Prag.
1 7 1 1 .R u r n p h i u s ,G . E , . . T h e s a u r u s c o c h l e a r u m . Lugduni Batavorum.
1 9 1 2 . P o m p e c k j ,F . J . , C e p h a l o p o d a( P a l i i o n t o l o g i e ) . Handu,cirterb.d. Naturw., Vol. II. Jena. G. Fischer.
1904. Sacco,F., i Molluschi dei terreni terziarii del Piemontee della Liguria. ParleXXX. Torrno. 1867. Schloenbach, U., Uber einenBelemnitenausder
1836. Quenstedt,F.A., Uber einige Hauptorganeder Nautileen.Arch. f. Naturf-.
alpinen Kreide von Griinbach bei WienerNeustadt.Jahrb.geol.Reichsanst., Vo1.XVII.
1836.Ibid., De notis Nautilearuntprintariis. Drss. Berolina.
1867-68.Ober Belentnitesntgi.ferSchloenb.Nov. Sp. aus dem eocrinenTuffe von Ronca. Ibid. Vol.
1839. Ibid.,Loligo Bollensisist kein Belemnitenorgan. N. Jahrb.f. Min.
XVll and XVIII. 1868. Ibid., lJberSepia Vindobonensis Schloenb.Nov.
1843. Ibid..Die Flozgebirge Wtirttembergs. Tribingen.
Sp. aus dem neogenenTegel von Baden bei
1 8 4 6 - 4 9 .I b i d . , D i e C e p h a l o p o d e n(.1 . V o l . Z u r
Wien.Ibid. Vol. XIX.
Petrefaktenkunde Deutschlands. ) Ttibingen.
1820. Schlotheim"J.F.v., Die Petrefaktenkundeauf
1 8 5 2 . I b i d . , H a n d b u c h d e r P e t r e l a k t e n k u n d e(.3 r d edition.I 885.)Tribingen.
ihremjetzigen Standpunkte.Gotha. 1913. Schwetzofi,A.S:, Les B6lemnitesinfracr6tac6es
1858. Ibid.,Der Jura.Tiibingen.
de I'Abbehasie.Annuaire 96o1.min. Russie,v.
1829. Rang, M.S., Manuel de 1'histoirenaturelledes
II. (15) Juriew. 1865. Seely,cf-.Rep.Brit. Ass.Adv. Sc.(p 100).
Mollusqueset de leur coquilles.Paris. 1 8 2 9 . R a s p a i l ,F . , H i s t o i r en a t u r e l l ed e s B d l e m n r t e s . Ann. Sc.d'Observation. Paris.
London.
1893. Renevier,E., Bdlemnitesaptiennes. Bull. Vaud., Sc.N., V. XXIX Lausanne. 1 8 5 4 .R e u B , A . E . ,
1913. Sinzorv.J.. Beitriige zur Kenntnis der unteren Kreideablagerungen des Nordkaukasus.
Loliginidenreste in
der
Kreideformation.Abh. k. bcjhm.Ges.Wiss. (5).
(Travauxdu Musde Geol. Pierre le Grand.Acad. Imp. Sc.Pdtersbourg, t. 7.) 1855. Sismonda.A.. Sur la constitutiongeologiquede
Vol. VIII. 1886.Riefstahl, E., Die Sepiaschaleund ihre Beziehung
1855. Simpson,M.. The fbssilsof the YorkshireLias.
zu
den
Belemniten.
Vo1.XXXII. Palaeontographica, 1856. Romer. F.. Uber Paloeoteuthis Dunen.si.saus d e m d e v o n i s c h e n G r a u w a c k e s a n d s t e l \nr o n
la Tarantaise ... C. R. Paris. 1916. Smith, E.A., On the shellsof South-African Speciesol the Sepiidae.Proc.Mal. Soc.,v. XIl. London. 1 8 2 9 . S o w e r b y ,J . d e . T h e M i n e r a l C o n c h o l o g yo f
Daun a. d. Eiiel (ein devonischerSepiaknochen
greatBritain, v. VI. London. (Deutschbearbeitet
...) . N. Jahrb. f. Min. cf.: Palaeontographica,
v o n E d . D e s o r . D u r c h g e s e h e nu n d r n i t
Vol. IV. Taf. 13:Lethaeageogn..Vol., I, p. 520.
Anmerkungenund Berichtigungenversehenvon
1858. Ibid., Zweites Exemplar von Archaeoteuthis Dunensis aus dem Tonschietievon Wassenacl.r am LaacherSee.N. Jahrb.f. Min. 1 8 9 0 .l b i d . . P l a g i o t e u t h i s , e i n e n e u e G a t t u n g dibranchiaterCephalopoden aus dem russ.Jura. Zeitschr.d. D. geol.Ges.,Vol. XLII. 1 9 1 0 . R o l l i e r . L . . M a t e r i a u xp . l a c a r t eg e o l . s u i s s e . (Nouv.56r.Livr. 25 [3. suppl.]) 1 9 0 8 . R o m a n , F . . R n i s i o n d e q u e l q r - r eess p d c e sd e
Dr. L. Agassiz. Solothurn1842.) 1910. Sprengler,E., Die Nautiliden und Belernniten desTrichinopoly-Distrikts. Beitr. z. Pal. u. Geol. Ost.-Ung.u. d. Orients,Vol. XXIII. 1 8 9 9 . S t e i n m a n n ,G . . U b e r d i e B i l d u n g s w e i s ed e s dunklen Pigments bei den Mollusken, nebst Bemerkungen tiber die Entstehung von Kalkkarbonat.Ber. naturf. Ges. Freiburg i. B., Vol. XL
Bdlemnitesdu julassicluerroven du jard et de
1903. Ibid.,Einftihrungin die Paliiontologie. Leipzig.
I'arddche.Bull. Soc.Sc.nat.Nirnes.v. XXXVI.
1910. Ibid.,Zur PhylogeniederBelemnoidea. Zeitschr.
1 8 2 9 . R t i p p e l l , E . . A b b i l d u n g c n L r n dB e s c h r e i b u n g einiger neuerl und u cniser gekannten Versteinerungen von Sohrhofin.Frankfurta.M.
f. induct. Abstammungu. Vererbungslehre,Vol. I\/
1890. Steinmann,G. u. Dciderlein,L.. Eler.nente der
t94 prisca Mrinst. 1835.Ibid., Notice sur 1' Onychoteuthis
Paliiontologie . Leipzig. 1 8 8 1 . S t e e n s t r u p ,J . , P r o f . A . E . V e r r i l l s t o n y e
et sur les Loligo Bollensiset Aalensis Zteten.
Cephalopodslagter: Stenoteuthisog Lestoteuthis.
L'Institut,journ. gdn. soc. trav. sc. Fr. 6tr., No.
Overs.DanskeVid. SelskForh.
147,Paris. Versuch einer
1 8 3 6 . I b i d . , N o t i c e s u r l e s r a p p o r t sd e s B d l e m n i t e s
geognostisch-botanischen Darstellungder Flora
avec d'autescoquillesintemesde Cephalopodes.
der Vor.welt.Letpzig u. Prag.
I b i d . N o . 1 5 7 .( c f . N o . 1 9 0 u . 1 9 6 . ,a n d c f . N .
1820. Sternberg, K.
Graf
v.,
1863-65.Stolickau. Blanford,FossilCephalopda of the Cretaceous Rocks of Southern India. (Pal. India.) Mem. Geol. Surv.EastIndia. Calcutta. 1 9 1 1 .S t o l l e y , E . , B e i t r l i g e z v r K e n n t n i s d e r C e p h a l o p o d e n d e r n o r d d e u t s c h e nu n t e r e n Kreide.Geol.Pal.Abh. (N. F.), Vol. X. 1911. Ibid., Studienan Belemnitender unterenKreide
J a h r b .f . M i n . , p . 1 8 5 , 3 2 3 u . 6 2 9 , a n d 1 8 3 7 p.723.) 1840. Ibid., Observationssur les Belopeltis ou lames Inst.t. 3. Paris. dorsalesdesBdlemnites. 1860. Wagner, A.. Die fossilenUberrestevon nackten Tintenfrschenaus dem lithographischenSchiefer Juragebirges. und dem Lias des stiddeutschen
Norddeutschlands.4. Jahresber.niedersiichs.
Abh. Mat.-phys.Kl. Kgl. Bayer. Akad. Wiss.,
geol. Ver. Hannover.
Vol. VIII.
1919.Ibid., Die Hiboliten und Neohiboliten der
1 8 3 2 .W a g n e r , R . , A c a n t h o t e u l i r l . s ,e i n n e u e s
unterenKreide in der Lethea geognostica.12.
Geschlechtder Cephalopoden,zu der Familie
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gehorend.Miinster, Beitriige,1. 1893-94.Walther,J., Einleitungin die Geologie.Jena, G. Fischer.
1 8 6 5 .S u e B , E d . , U b e r d i e C e p h a l o p o d e n s i p p e Acanthoteutlis. Sitz.-Ber.Wiener Akad., Vol.
1904. lbid., Die Fauna der SolnhofenerPlattenkalke, bionomischbetrachtet.Festschr.f. Haeckel. 1911. Wanner, J. Triascephalopoden von Timor und
LI. 1 7 3 8 . S w a m m e r d a m ,J . , B i b l i a n a t u r a e . V o l . I L
Rotti. N. Jahrb.f. Min. Suppl.Vol. XXXII. 1 9 0 9 .W e i g n e r , S t . , S t u d i e n i m
Leyden. 1893. Tate, R., Unrecordedgenera.Journ.Proc. Roy. Soc.New SouthWales,v. XXVI.
Gebiete der
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1885. Teller, F., Fossilfiihrende Horizonte in der
1 9 1 2 .W e r n e r , E . , U b e r d i e B e l e m n i t e n d e s
oberen Trias der SanntalerAlpen. Verh. k. k.
schwiibischen Lias und die mit
geol.Reichsanst.
verwandtenFormen des braunen Jura (Acoeli).
1909.Till, A., Uber fossile Cephalopodengebisse. Verh. zool.-botan.Ges.Wien, Vol. LIX. 1900. Tornquist, A., Die Arbeiten der drei letzlen Jahre iiber
fossile
Cephalopoden. 5.
Triascephalopoden.Z. Zentralbl., J ahrg. 7 . 1 8 7 9 .T y r o n , G . W . , M a n u a l o f
Conchology.
Philadelphia. 1882. Tullberg, Studien iiber den Bau und das Wachstum des Hummerpanzers und der Molluskenschalen. Kgl. Sv. Vet. Akad. Handl., Vol. XIX. 1 8 8 1 . V a s s e u r ,G . , R e c h e r c h e sg 6 o l o g i q u e ss u r l e s t e r r a i n s t e r t i a i r e sd e l a F r a n c e o c c i d e n t a l e . Stratigraphie.Parrs. 1830. Voltz, P.L., Observationssur les Bdlemnites. Mdrn. Soc.Mus. Hist. N. Strasbourg,t. L Paris.
ihnen
Palaeontographica, Vol. LIX. 1889.Whiteaves, Contributionsto the Canadian Palaeontology, v. I. 1902. Willey, A., Contributionsto the naturai history of the pearly Nautilus. Zool. Results Willey. Cambridge. 1863. Winkler, T.C., Catalogue systematiquede la C o l l e c t i o n p a e o n t o l o g i q u ed u M u s . T e y l e r . Harlem. 1903. Wojcik, K., Die unteroligociine Fauna von Krihel-Maly bei Przemysl. I. Bull. Acad. Sc. Cracovie.(1904?) 1851.Woodward, S.P., Manual of the Mollusca. Suppl. 1856.(new editions) 1883.Woodward,H., On a new Genus of Fossil ,,Calamary",from the CretaceousFormationof
19s Sahel-Ahna,near Beirut. Libanon, Syria. Geol.
Abbildungen und Fundorte. Corr.-Bl. Landw..
Mag. (2) v. X.
Ver.,Vol. I.
1 8 9 6 . I b i d . . O n a F o s s i lO c t o p u s( C a l a | s N e v t b o l d i ) frorn the Cretaceousol the Libanon. Quart. J. Geol.Soc.,v. LII. London. 1896. Ibid., Calai's llev,boldi (preoccupied).Geol. M a g . ,r ' . I I I . 1901. Yoshiwara,S., On an apparentlyner''.species of
1868.Zittel.
K.A.v.,
Die
Cephalopoden der
StrambergerSchichten.Pal. Mitt. Mus. Kgl. bayr.Staates, Vol. II. 1885. Ibid., Handbuch der Palaeontologie,Vol. II. M i i n c h e nu . L e i p z i g( l 8 S l - 8 5 ) . 1895. Ibid.. Grr-rndztige der Palaeontologie. Mtinchen
Argonauta from the Tertiary of Izumo. Annot.
u. Leipzig. Neubearbeitetvon F. Broili. 2.
Zool. Jap.,v. Iil.
e d i t o n1. 9 0 3 , 31. 9 1 0 , 41. 9 1 5 . 51. 9 2 1 .
1822. Young, G., u. Bird. J.. Geologyof the Yorkshire c o a s t2. . e d .W i t b y 1 8 2 9( 1 8 2 8 ) . 1830. Zieten, C.H.v., Versteinerungen Wtirttembergs. Stuttgaft. 1839. Ibid., GeognostischesVerzeichnis siimtlicher Petrefakten Wiirttembergs mit Zitaten. ihre
Supplement
(Some literaturereferencesare given in the text. A discussionof oldervierr,,s oppositeto oursis not alu.ays explicitly given. Whereverour descriptiondiffers from older ones,such a discussionshouldbe consideredas tacitlyassumed).
s c i e n c e sA. n 1 . S c . N a t ( 2 ) t . 4 . Z o o l o g i e .( c f a u c hC . R .P a r i s1 8 3 5p. . 3 a 1 . )
1 8 3 8 . B e l l a r d i ,L . . N o t i c e s u r l ' A r g o n a u t an i t i d o L k . B u l l . S o c .G e o l .F r . ( 1 ) t . 9 . P a r i s . 1880. Bukman,J.. On the Belemnoteuthis Montefiorei. Proc. Dorset.N. H. antiquarianField Club v. 3 (1879). 1 8 4 7 . C u n n i n g t o n .W . . O n t h e f o s s i l C e p h a l o p o d a lrorn the Oxford-Clay consistutingthe genus Belemnoteutlri.s Pearce.London, Geol. Journ. R e c .D i s c o v e r i e B s r i t . f o r e i g nP a l e o n t o l o g yr ., . l' 1824. Cuvier, G., Sur des os de Sechefossiles.Ann. S c .N a t . ,t . 2 . P a r i s . 1848. Dana, J.D., Fossilsof the exploringexpeditiolt u n d e r t h e c o m m a n do f C h a r l e sW i l k e s . . . a Belemnitefiom Tierra del Fuego.SillimanAm. Joum.Sc.Arts. (2), v. 10. 1864. Dawkins,W.B.. On the Rhaeticbedsand White L i a s o f W e s t e r n a n d C e l r t r a l S o m e r s e t .Q u . Journ.Geo1.Soc.London,v. 20. 1866. Deshayes.G.B.. Descriptiondes animaux sans vertebresddcouverts dansle bassinde Paris,v. 2
1 9 1 1 . F r . i c ,A . . S t u d i e ni m G e b i e t ed e r b c i h m i s c h e n K r e i d e f o r m a t i o nA . rch. f. nat. Landesdurch_ forschungvor1Bohnel. Vol. 15.prag. 1861. Gabb. W.M.. Dcscription of a neu, speciesof C c p h a l o p o df.i ' o m t h e E o c e n eo f T e x a s .P r o c A c . n a t .S c .p h i l a d e l p h i(a1 g 6 0 ) . 1883. Guettard.Sur.les bdlernnites. Mdm. difl-.oarties paris. Sc.Arts. t. 5. 1859.Hehl. Die geognostischen Verh6ltnisse yu'tirttembergs. (Syst. Verz. der Belemniten.p. 206.)Stuttgart. 1 8 8 9 . I s s e l . ,A . D i u n a S e p i ad e l p l i o c e n ep i a c e n t i r . r o . Bull. Soc.mal. Ital. v. 14.Roma. 1 8 4 5 .K u r r ,
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1 8 7 6 . T a t e , R . , & B l a k e , J . F . , T h e Y o r k s h i r eL i a s . London. 1861. Trautschold,A., Recherchesg6ologiquesaux environs de Moscou. Bull.
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1 ' d o c d n eb e l g e . C d p h a l o p o d e sd i b r a n c h i a u x .
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of the fossil 1856. Woodward,S.P.,On the Occurrence g e n u s C o n o t e u t h i s d ' O r b . i n E n g l a n d .A n n . Mag. Nat. Hist. (2) v. 17.London. 1829. Young, G., & Bird, J., A geologicalsurvey of the Yorkshirecoast.2. ed.
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Furtherliteraturecan be found in v. Brilow 1920,
r9l
(317)
alveolarslits 200, 201x,241.
Index with notes
r.)
Amhlvbeltrs205x, 279, 291. Ancistroteuthls I 89x, 298.
a a l e n s i s( B e l o p e l t i s . . . ,L o l i g o . . . , B e l e m n i t e s . . .1) 0 5 , 1 0 9 ,1 2 5 ,1 2 9 x , 1 6 9 . Abel. O. (fundamentalremarks)7, 165,214,219,221, 228.
apicalfunows 198. apicalline 204x,206. Architetrthis101,298. Argonauta287,288x, 291x,293x,300,frontispiece.
Ac'anthopus 175.
arm number29, 160,219,252x.
Ac'anthoteuth is 29, 115, 177.
Ascoceras95.
A. conocattda172x,179,189x.
Asteroconites267.
A . j o e c k e l i1 1 2 x , 1 1 9 .
a s y m p t o t e3s 1 x , 3 2 , 1 0 5 , 1 2 3 x , 1 2 9 x . 1 4 8 x , 1 7 0 x ,
A . m o n t e . / i o r e1i 1 6 ( s e e a l s o C r i c k 1 9 0 7 : A r m s o f 178,I 89x. .,belemr.rites")
2 0 q x . 2 1l x . 2 5 0 x . "asymptotefurrows" 265.
A. prisca 175.
Ah'actites263x,270, 216, 281.296.
A . p r o b l e n t a t i c1a5 l x , 1 5 2 ,1 8 3 .1 8 9 x .
The genuscomprisesboth the oldest and the youngest
A. speciosa 1 1 0 x , 1 1 4 , 1 8 01, 8 9 x ,2 5 0 x , 2 5 2 x .
aulacoceratidsand representsthe main stem of
accretion lines (striation), very important for the
the older belemnoids,also in terms of their
reconstructionof shell fragments(outline and
diversity. It certainly has to be subdividedin
curvature)and their development:110x, 123x,
groups the rank and relationshipsof which can
1 2 9 x .1 4 3 x ,1 4 8 x . 2 0 4 x , 2 0 9 x . 2 5 0 xT.h e r ea r e
not yet be determined with certainty. In any
threesoftsof striation:
event, the following types have to be
1.
2.
marginal striation due to the step-wise
distinguished:
enlargementof the shell surface, in general
a)
(but not always) arrangedin concentricline
club-shaped in lateralvieu'. Distinct,widely
patterns or excentricaliy progressingparallel
spaced lateral furrows. similar to
to the margin (1a3x). Inside the nucleuslies
Calliconites dieneri (see there), to which
the primordial shell.
they probably belong. Here belong: l.
longitudinal striation due to the shifting of
porr:Lrs, gracilis, sundaicus, aclrtus,
certain points of the shell epithelium (matrix)
lanceolatus, v. Biilow 1915.
radiatingfrom the primordial shell (137x); if
3.
rostra stronglycompressedlaterally,slightly
b)
rostra moderatelycompressed,elliptical in
the initial part is missing, these lines point
cross section, without distinct lateral
towardsthe primordialshellposition(123x).
furows. Here belongs:A. cylindricusHauer
striation from stratification at fracture or
( v . B i i l o w 1 9 1 5 p, . 5 8 ) .
sectionpoints (like 1) (20ax). {Patternsand sculptures are often due to uneven surfaces
c)
rostrabarely or not at all compressed, subquadraticto nearly circular in cross section,
corresponding to growth lines. However,
rvithout distinct lateral furrows, ventrally
secondaryincrustationscan run across the
u i t h s l i g h td e p r e s s i o n . c)
primary growth lines (ll0x) so that uneven
cylindrical to conical, short, in
surface parts can be adopted from different
lateral view. Alveolus strongly
matrixpositions(288xd) | .
excentrical.Here belongs:l.
Actinocamax256x.291.
quadratoides
A Iloteuthi s | 65.211x, 220.297.
( 1 9 1 0p, . 1l 8 ) .
A l v e o l u s1 7 0 x( F i g .6 3 d ) ,1 7 5 ,1 9 5 .
ti)
Steinmann
stocky, club-shapedin lateral
alveolite195.
view. Here belongs: A.
alveolarfurrows 200, 201x, 246x,211 (ventralgutter, "canal").2.)
c'laviger,Biilow I 9 I 5. d)
Rostra slenderand club-shaped.circular in
198 c r o s s s e c t i o n , s i m i l a r t o H a s t i t e s .H e r e
burying in Sepioidea43. Typical octopodidsshow a
Hauer 1887.Karn.belongs:A. terutirostrl.s
similar behavior after the end of the pelagic
norian level, perhapsa precursorofhastites.
larval condition (cf. Cephalopoda,vol. I, chapter
AtrIacotetrthi s 243x, 245,246x,291.
50).
Aulacoceras 263x,267,296. 262,263x,296. Aulacoceratidae
Calai's285.
axial thread203,204x, 206.
calamary(squid)101. caicareous cones14x, 16. C a l l i c o n i t e s 2 7 3 , 2 9 6 ( i s n o t a p r e c u r s o ro f t h e
259, 291. Bayanoteuthinae
Teuthoideaas supposedp. 104 and 162).
Bayanoteuthis259x, 291. BelemniteIla 194,201x, 255, 297.
capitulum45x,62x,63x, 64x, 66x, 76x, 77x,83x.
Belemnites, generalsignificance194, typical fossils
cartilaginousrods 285.
I 70x.
Celaenidae150,29'7.
(Hibolites)168,113,180,250x,251. B. semisulcatus
181,184),291. Celaeno150, 151x(erroneously:
Belemnitidae 193, 194,221, 224,296.
753x,291. Celaenotetrthis
arm nunrber(see there), development204x, 209x, life
Cephalopoda. Shell type 14, Diagnosis17, Systeml8
style 220, phragmocone(see there), pro-
( s e ea l s oN a e f 1 9 2 1 p, . 3 1 0 ) .
ostracum(seethere), reconstruction(see there),
chamberedsnail24.
rostmm (seethere). 246x, 291. Belemnoconus
27,298. Chaunoteuthis Chiroteuthis217,298.
B e l e m n o i d e3a0 , 1 6 4 ,1 6 6 x , 2 9 6 .
Chiroteuthidae219.
247,291. Belemnopsinae
Chirothauma165,218x, 298.
Belemnopsis246x, 249, 297.
Chitinousrings or "horny rings": cuticular structures
Belemnosella 31x, 33, 49x,291.
supportingthe suction chamberwall in decapod
109,169,ll 3. Belemnosepia
suckers(cf. p. 27,Fig. 5b: x, y, z) 28.
48,299. Belemnosidae
101. Chondrophora
Belemnosis 49x, 50, 51,x,299.
24. chromatophores
Belemnoteuthidae 276, 297.
Cirroteuthoidea 284, 285, 300.
B e l e m n o t e u t h i sl l 3 ,
Clavirostridae207, 228.
186x, 189x, 276, 297,
erroneously:172x.
club-shapedrostrum192,220, 226.
125,291. Belopeltidae
2 2 9 ,2 9 6 . C o e l o t e u t h i en a
Belopeltis125, 129x, 169, l1 | , 291.
Coeloteuthis229, 23lx, 296.
Beloptera55, 56x, 51x, 299.
conchiolin12.
Belopteridae 53,299.
c o n c h i o l icno n e sl 4 x , 1 6 .
Belopterella 59x.
Conirostridae201, 228.
Belopteriditm 53x, 54, 299.
251x, 258,291. Conobelus
Belopterina 53x, 54, 299.
conotheca15.
Belosepia39x, 80x, 82, 83x, 299.
Conoteuthis112x,278, 297.
Belosepiella 60, 280x, 299.
c o n u s 1 0 3 x , 1 0 5 , 1 5 8 , 1 5 9 x , 1 6 0 x ,2 1 7 x ( r e m a i n so f
Belosepiinae 81,299.
chamberformation 157x, 2 18x).
Beloteuthidael4l, 291.
conusvane 105,108.
Beloteuthis142, 143x, 148x,29'7.
corrosionforms201,256x.
Benltteuthis(Gonatidae)29, 298.
Cranchiidae299.
bollensis(Loligo, Beloteuthis,Geoteuthis)125, 144,
Crassicarinati 135.
165. bow region 108: middle plate. Brachybeltts204x, 23 lx, 233x, 241, 296. Biilow, E. v. 223,263,266, 305.
Ctenoglossa 300. curvatureof apical line compensatingventral curvature of pro-ostracumand phragmocone47,210. Cylindroteuthinae242, 297.
r99 C.vI indroteuthis 242x,243x,291.
G o n , t t r r1s5 7 x .1 5 9 2. q 8 . "Gonioteuthis"256x.
Dactyloteuthrs204x, 231x.236, 244,291. D e c a p o d2 a5 , 2 9 6 .
growth of molluscan shell 13; cf. growth lines: accretion.
developmentalnorms, morphologicainorms etc.: see norms.
"Hastatidae"247.
developmentof the belemniterostrum204x.
Hastites222x, 22 6, 221x, 296,
Dibranchiata19,21 (Diagnosis), 24, 296.
Hastitinae225,296.
dibranchiatecephalopod21, 24.
head-foot11.
Dicoelites254,291.
"Heliceras" 68.
D ict.voc' oni tes263x, 270. 296.
Hemisepius 84,299.
Diploconus33, ll2x, 278,291.
Heteroglossa 300.
dorsalshield78 ; layers91x.
Hibol ites204x, 249, 250x. 291.
doublefurrows 198.
Homaloteuthis204x. 241,296.
"Dorateuthis"l18.
h o o k s2 6 , 1 8 7 "1 8 9 x , 2 5 2 x .
"Doratopsis"217x.
horny ring (cf. chitinousring).
"Dorvanthes"7l5.
hump (of cuttlebone)79.
Dtrvalia257x,258,291.
hydrostaticsof phragn-rocone 17.
Duvaliinae257,291.
hyperboliclines I 08. 174,204x,209x (cf. lateralplate 211x).
Eledone286x,300.
hyperboliczone(:lateral plate 21 1x).
elongation(secondaryelongationof phragmocone) 47.
hypostracun.r 12x,13, 14x, 15, 107(cf. nacreouslayer).
5lx, 64x,73. embryosof Loligo 159x,Sepia98x, Sepietta98x.
Idiosepius33, 91. 299.
"embryonicrostrum" (so-called)accordingto Stolley
Ill er 220. 298.
204x, 206-208;correctterm: juvenile rostrum;
i n k s a c2 4 , l l 7 , 1 9 5 .
cf. p. 268. "embryonicchamber"(so-called);correctterm: initial
juvenile rostlxm 204x (cf. embryonicrostrurn).
chamber207. embryonic shell of belemnites 204x, 209x, of Sepia 39x,91x, 98x,of Spirula73.
Kelaeno(cf. Celaeno). Kelaenidae(cf. Celaenidae).
"embryonicthread"203; cf. primordial rostrum. E n d o c e r a t i d a1e6 , 9 5 .
lateralbow zone 105,2l lx.
Enoploteuthis298 (enoneously:I l6).
lateraifield of middle plate 121 (cf. niiddle field).
excentricityof rostrumgrowth 41, 210.
l a t e r afl u n o w s 1 9 8 "2 6 5 . laterallinesofconotheca21 1x.
fins: development36, relation to muscular mantle and shell34x, I l4x, l60x (a2)
lateralwings 51x,64x. lateraledges54x, 62x (in 3).
Fork 77x, 79, 80x, 86x.
lateralbulges44x, 5 1x.
funnel apparatus22.
L e a c h i a2 1, 2 9 9 .
funnel attachment26.
Leptoteuthis ll9, 1,20x,291 .
funnelfunction23.
l i f e s t y l e : S e p i o i d e a4 3 , T e u t h o i d e a1 0 4 , 1 5 6 , 1 9 3 .
funnelvalve 26.
B e l e m n o i d e1a9 1 - 1 9 3 . Liogiossa300.
Gastroheltrs 233x, 234. 296.
Lioteuthi.sl l0x, 132, 291.
122,123x,l24x ("G. :itreli" l33x), 297. Geotettthis
Listroteuthis193.
g l a d i u s3 2 , 1 0 5 ,1 5 7 x .
"lobe ofthe ear" 161.
Gl.vphiteuthis 134x,1 40, 297.
Loliginites102.129x,130,291.
200 Loligo 155x,297;L. sagittata116,190.
Palaeoctopoda 18, 285,300.
"Loligosepia" 122, 125, 126,252.
Palaeoctopus 285,286x,300. 147, 148x. Palaeololiginidae
mantle12x,31,32.
125. Palaeo.sepia
mantle sac I I (i.e. the body part typically enclosedin
paleomorphology2, 8, 296.
the shell of a mollusk, in contrastto the head-
Parabelopeltls108,128, 129x,297.
foot; cf. Naef 191I, p. 83)
parabolarfield 31, 105, 108,21lx (rniddleplateof pro-
Megatetrthis 204,237x, 239, 296.
ostracum).
Mesohibolites 254, 29'7.
Parahibolites254,291.
Mesoteuthoidea 32, 135,291.
Paraplesiotettthis110x, lll,
Metasepia 84, 299.
Passaloteuthinae 230, 296.
Metateuthoidea 32, 154,291.
Passaloteuthis204x, 232, 233x, 296.
middleplate31x,32,105,209x,210,211x,250x. middle field 108 (here : middle plate), 114x, 121 (pafi
periostracum 12x, 13,44, l0'7, 176.
of the middle plate).
129,297.
Philpott168. phragmocone 15,212.
Mollusca,schematicrepresentation12x.
Phragmoteuthis ll2x, 186x,189x,261,296.
m o r p h o l o g y , b a s i c c o n s i d e r a t i o n s2 , 4 , 2 9 6 , c f .
P hylloteuthis149,297.
paleomorphology.
phylogen2 y ,3,296.
muscularmantle22, 24, 31,42x, 191.
phylogenyas pseudogenealogy 2-6.
Myopsida19, 40, 101, 191,297.
p i l l a r sl 4 x . | 5 . l 3 x . 7 4 x . P l e s i o t e u t h i d|a|el , 2 9 1.
nacreouslayer 13, 240 (cf. hypostracum).
Plesioteuthis 110x,113, ll4x, 297.
Naef, A. 310 (see the Foreword about the unity of
Pletrrobelus233x, 235.
working plan).
284,300. Polypodoidea
Nannobeltrs231x, 232, 297.
"Polyteuthidae"201.
Neohibolites254x, 297.
primarylid 21.
norms (developmentalnorm, constructionalnorm), 6,
primary mantle(:dermal mantle) 12x,22.
8-10,296. nuchal attachment22.
primordial rostmm 203,204x. pro-ostracum 3 1x,209x,210,21 lx. "Protodecapus"I67x.
Octopoda25, 283,300.
"Protosepioides"45x.
25, 285,300. Octopodidae
"Prototeuthis" 103x.
27, 298. Octopodoteuthidae
Prototeuthoidea 32, 108,297.
286x.292.300. Ocroptr.s
p s e u d o a l v e o l2u0s 2 .2 5 6 x .
O c y t h o €2 9 1 , 3 0 0 .
Pseudobelus 202,257x,291.
Odontobelus 204x, 23 lx, 238, 296.
Psettdodttvali a 258, 297.
O e g o p s i d a 4 01,0 1 ,1 5 8 .1 6 0 x ,1 9 1 , 2 9 8 .
Pseudohas tites 234,243, 296.
"Ommastrephes"I15, 116,O. meltrati 135.
Ptiloteuthis150,291.
Ommatostrephes I l, 298. 220, 298. Ommatostrephidae
Q u e n s t e dFt ,. A . 1 0 2 ,1 0 8 ,1 6 9 ,1 7 3 , 2 0 3 , 2 2 1 .
"Orychites"187,189x. Onl,choteuthisI 60x, 298.
radula39.
" O . p r i s c a " 1 0 2 ,11 5 , 1 1 6 , 1 2 5 , 1 6 81, 7 8 ,1 9 0 .
reconstruction, basic principle 7, belemniteshell 168,
281. orthoceres
2 0 4 x , 2 0 9 x , 2 1 1 x2, 5 0 x ,b e l e m n i t ea n i m a l1 8 6 x ,
outerplate (middle,inner plate,cf. dorsalshield).
204x, 209x, 211x, 213. Earliertentatives:214x
Oxyteuthis204x, 243x, 244,291.
275x, 276x. Abel: 220x. Other forms: 54x, 57x, 1 2 0 x ,1 3 9 x ,1 8 6 .S h e l l s 4: 9 , 5 1 , 5 9 ,6 2 , 6 4 , 1 6 ,
Pachltetnhis204x, 244, 297.
| 11, 120, 123, 124, 129, 137, 250.
201 regulationof longitudinalaxis 47, 2 I 0; seealso:ventral curvature(of phragmocone).
staticsof air chambersI 9 I . Steinrnann, G.9,271,213.
Rhabdobeltrs 228,251,296.
Sthenoteuthis 157x,165, 298.
Rhaphibelus 245.250x,291.
Stolley,E. 206,223,228.
"Rhopalohelus" 226.
Stratificationof shells 105-107.109 (gladii and pro-
"Rhopaloteuthis"258.
ostraca).
Rondeletiola99,299.
Styloteuthis119,297.
rostrum, in strict and broad sense: l7 5-116:
St,vracoteuthis 259,260,291.
development203, 201x, 268x; function 192;
suckers2T.
main forms 197x.
supporlingridges 15. ing movements(cf. funnel function).
Salpingotettthis 235,231x, 243,246x, 296. Sepia,juvenile form: 39x, 78x, 98x; embryonicshell:
tentaculararms25x, 167x.189,252x.
39x, 91x, 98x; shell structure80x. 86x, 88x.
"Tenuicarinati"135.
91x; fossil forms: 92. spurious:93; systematic
"Tenr-rimargir.rati" 208.
position299.
Tetrabranchiata 19,20.
"Sepia"venusta291.
"Teudopsis"130,136.190.
"Sepialites"130,134x.
"Teuthidae"173.
Sepiidae78.299.
"Teuthis"32.
Sepiinae84,299.
"teuthods"181.
Sepioidea32, 38, 45x,299: development39x; life style
Teuthoidea 32. 101. 103x,291.
43.
"Teuthopsis"147.
Sepiola33,299 (seealsoSepietta).
"Tettthos"32.
Sepiolidae 91,299.
theoryof derivation9^ 10,296.
"Sepiophora"94.
theoryof derivationand morphology4, 6. 9, 10.
"Sepioteuthis" 102,169(in the correctsense:297).
Tht'sanotetrthl.s 261,298.
shell (cuttlebone)3 I (types)
Trachyteuthida e 136.291.
shell grorvtlr,seeaccretion.
Trachyteuthis134x, 137x, 139x,291.
shellsac24.
Trentoctopus 291,300.
shell sculpture.seeaccretion.
Tschulok,S. Theory ofheredity, Jena1922.
siphunclel5 (alwaysventral, marginal in dibranchiates
Tusoteuthis149,297.
264). speciesconcept6.
Varxpyroteuthidae 285, 300.
Spinrla 69, 299; juvenile form 39x, 41x, 73x; adult
vane(cf. conusvane)107-108.
6 9 x , 7 1 x l,3 x , l 4 x .
Vassetrria259x, 280, 297.
Spirulidae68,299.
Vasseuriidae280,297.
Spirtrliro.stra39,62,63x.64x,66x,299.
ventralcurvarureofphragmocone4T.
Spirulirostrella50x,299.
venrralrim 40.
Spirulirostridae 60,299.
ventralwall 33. 44,105.
SpirttI irostridi um 61, 62x, 299. Spit'ttlirostrina 75,16x,299.
Werner223.
"Spirulisepia"77x. stabilization of equilibrium95.
Xiphoteuthis113x,275,296.
202 List of new species:
t.)
Bold-facednumeralsindicatethe relevant foriginal]
page numbers, crosses(x) indicate the figures on a l.
jaeckeli 172x,179. Acanthoteuthis
given page. As to the position of the families, see
2.
problematica151x,183. Acanthoteuthis
Figurel0l (p. 303).
3.
Belemnosiscoszmanni5lx.
1.
Belemnoteuthisaclttalllx,246.
'.;
S.. Neumayr, M.: On some belemnites from
I propose this designation for the slender
CentralAsia and on the canal of belemnites.Verh.
phragmoconeswith sheathshown in Fig. 64c,
Geol. Reichsanst.Wien. 1889.
which cannotsafelybe placedelsewhere. 5.
Belopteralonga 56x,57x.
6.
Belopteridiumpuerilis 110x, 132.
7.
incerta 153x. Celaenoteuthis
8.
Lioteuthisproblematica110x,132.
9.
Paraplesioteuthis magnall0x, l12.
10.
Spirulirostrasepioidea66x,6T.
11.
Spinrlirostridium obtusum61,62x.
See p. 296-300 on new names and establishmentof generaand families.
203
Appendix
B a t h e r i n B l a k e , 1 8 9 2 , B e l e m n o i d e a )( n o w strongly restrictedto speciesclose to the type species A. alpinusGiimbel,1861).
List of synonyms
Callic'onites Gemmellaro, 1904 (valid genus of
by Theo Engeser
Belernnoidea.indet position). A . p a r v u sv . B t i l o w ,1 9 1 5 . A. gracilisv. Biilow, 1915.
aalensisZjeIen, 1832 (Belopeltis in Naef) (valid
A. sundaicusv. Btilow. 1915.
speciesof LoligosepiaQuenstedt, 1839).
A . a c u l t t sv . B r i l o w .1 9 1 5 .
A c a n t h o p u s M i i n s t e r , 1 8 3 9 ( j u n i o r , s u b j e c t ier
A. lanceolatu.r v. Btilow. 1915.
synonymy of Acanthoteuthis Wagner in
A. cylinclricus Hauer.1887.
Mtinster.1839).
A. cluadratoidesSteinmann,1910.
AcanthoteuthisWagnerin Miinster, 1839 (valid genus ol
Belemnotheutididae Zittel,
A.
1884.
Claviatt'actitesMariotti & Pignatti. 1996^family
Belemnoidea).
X i p h o t e u t h i d i d a eB a t h e r i n B 1 a k e " 1 8 9 2 ,
A. conocaudaQuenstedt,1849 (now usually assigned
Beiemnoidea).
t o P h r a g m o t e u t h i s M o j s i s o v i c s ,1 8 8 2 , b u t
A. tenuirostri.s Hauer,1887.
probablynew genus).
A u l a c o t e u t h l ^S r t o l l e y . 1 9 1 1 ( v a l i d g e n u so f f a r n i l y
A. jaeckeli nov. sp. (unior, subjectivesynonym of
Cylindroteuthididae Stolley,
P hr agmoteuth is ntonte/iorel Brickrnan,1880). A. ntontefiorelBuckrnan,1880 (now usually assigned probablynew genus). A. prisca - Plesioteuthisprisc'cr(Riippell, 1829).
1919.
Belemnoidea). Aulacocera.sHauer, 1860 (valid genus of farnily
to Phragmoteuthi,s Mojsisivics,1882 but
A.
c l a v i g e r v . B i i l o w , 1 9 1 5 ( t y p e s p e c i e so f
Aulacoceratidae Mojsisovics,
1882,
Belemnoidea). Ar-Llacoceratidae Mojsisovics, 1882 (valid lamily of
problematica n.sp.(indet. belemnoidcf.
Belemnoidea).
AcanthoteuthispeciosaMiinster, 1839 or Belemnotheutis mavri Engeser& Reitner,1981).
Bayanoteutl.rinaenov. (prob. valid farnily of the
A. speciosa Mtinster, 1839 (type species ol
Spirulida.not listed in Doy1e,Donovan &
Acanthoteuthr.s Wagnerin Mtinster. 1839). A c : t i n c t c o m a ,M r iller, 1826 (valid genusof lamily
Nixon. 1994). B a y a n o t e u t h l sM r . r n i e r - C h a 1 m a1sU. 7 1( v a l i d g e n u s ;
Belemnitellidae Pavlow.1914.Belemnoidea).
type genus of BayanoteuthididaeNaef. 1922,
AllotetrthisWulcker, 1920 (valid subgenusof Loligo Schneider,1784. Family Loliginidae Lesueur.
Spirulida). Belemnitella d'Orbigny, 1840 (type genusof farnily
1 8 2 1M . yopsida).
B e l e m n i t e l l i d a eP a v l o w i n S c h w e t z o f t 1 9 1 3 .
AmbyhelusNaef, 1922 (indet.belemnoid)( homonym of Amb lt:belusMontrouzier,1864).
Belemnoidea). BelemnitesLamarck, 1799(invalid taxon,seeRiegraf.
Ancistroteuthis Gray, 1849 (valid genus of Onychoteuthidae Gray, I 849). A r c h i t e t r t h i s S t e e n s t r u p ,l 8 5 7 ( t y p e g e n u s o f ArchiteuthidaePf-ef-feL, I 900). Argonauta Linn6, 1758 (type genusof Argonautidae Cantraine,1841). Ascoc'erasBarrande.l8-+r (a ralid genusof nautiloid cephalopod) Asteroc'onites Teller. 1885(.iunior.sub.jective synonym o f A u l a c o c e l a . rH a u e r . l S ( r 0 .t l d e M a r i o t t i & Pignatti,1999) AtractitesGtimbel,1861(ialid lrcnu: -\iphotcuthididae
Janssen & Schmidt-Riegraf, 1998). B.
s e n t i s u l c a t u sM i i n s t e r . 1 8 3 0 ( H i b o l i t h e s s e m i s u l c a t u s ( M r i n s t e r ,1 8 3 0 ) , f a m i l y Mesohibolitidae Nerodenko,
1983,
Belemnoidea). B e l e m n i t i d a ed ' O r b i g n y . 1 8 4 0 ( i n v a l i d t a x o n , s e e Riegraf,Janssen& Schmidt-Riegraf'. 1998). B e l e n t n o c o r ? r / .ns o v . g e n . ( r , a l i d g e n u s o f t a m i l y Mesohibolitidae Nerodenko,
1983,
Belemnoidea). B e l e m n o i d e a M a c G i l l i v r a y . 1 8 4 0 ( s u p e r o r d e ro f Coleoidea, sister-taxon of Neocoleoidea).
204 Belemnopsinaenov. subfarn.(invalid taxon, based on preoccupiedgenusBelemnopslsBayle, 1878). BelennopsisBayle, 1878 fiunior, primary homonym of BelemnopsisE,dwards,1849, older synonym of Lagonibelu.s Gustomesov,1958, family Cylindroteuthididae Stolley,
1919,
Belemnoidea).
of
T e u d o p s i s E u d e s - D e s l o n c h a m p s1,8 3 5 ,
family BeloteuthididaeWiltshire, 1869, Vampyromorpha). Bercyteuthis Naef, 1921 (valid genus of Gonatidae Hoyle, 1886,Oegopsida). BollensisZieten, 1832(Loligo) (subjectivesynonymof L oI igosep ia aaIensis Zreten, 1832).
Belentnosellanov. gen. (valid genusof Belemnoseidae Wiltshire,1869,Spirulida).
BrachybelusNaef, 1922 (unior, primary homonym of BrachybelusStil, 1869;replacedby Brevibelus
BelemnosepiaBuckland& Agassizin Buckland,1836 (invalid genus).
Doyle, 1992, family MegateuthididaeSachs& Nalnjaeva,I 967, Belemnoidea).
BelemnosidaeWiltshire, 1869 (valid family of the Spirulida,must be correctedto Belemnoseidae). BelemnoteuthidaeZittel, 1884 (valid family of the Belemnoidea, must
be
corrected to
Calai'sWoodward, 1896 fiunior, primary homonym of Calai's Rafinesque, l8l5;
replaced by
Palaeoctoptrs Woodward, 1896 and Calaita
Belemnotheutididae: basedon Belemnotheutis
S t r a n d , 1 9 2 8 , P a l a e o c t o p o d i d aDeo l l o , 1 9 1 2 ,
Pearce,1842).
Octopoda).
BelemnoteutlrlsPearce,1842 (valid genus;must be corrected, original spelling is Belemnotheutis Pearce.1842). Belopeltidae Naef, 1921 (invalid taxon, subjective
Calliconites Gemmellaro, 1904 (valid genus of Belemnoidea, inc. sedis). CelaenidaeNaef, 1922 (is based on the incorrect spelling Celaeno Owen, 1844 for Kelaeno
synonym of LoligosepiidaeVan Regteren
Mrinster, 1842,both spellingsare preoccupied,
Altena, 1949, Vampyromorpha).
replacedby Muensterella Schevill, 1950 and
Belopeltis Voltz, 1840 (unior, subjectivesynonym of LoligosepiaQuenstedt,I 839). Belopterade Blainville, 1827(valid taxon of the family BelopteridaeNaef, 1922, Spirulida;type genus of BelopteridaeNaef, 1922). Belopteridaenov. fam. (valid family of the Spirulida). Belopterella nov. gen. (valid genus of the family BelopteridaeNaef, I 922, Spirulida). Belopteridiunl noy. gen. (valid genus of the family BelopteridaeNaef, 1922, Spirulida). BelosepiaVoltz, 1830(originaland comectspellingis
MuensterellidaeRoger, I 952). Celaenoteuthis nov. gen. (valid taxon of the Muensterellidae Roger, 1952,Vampyromorpha). Chaunoteulftls Appeloi 1891 (valid genus of Onychoteuthidae Gray, I 849, Oegopsida). Chirotettthis d'Orbigny, 1841 (valid genus of ChiroteuthidaeGray, 1849, Oegopsida). Chiroteuthidae Gray, 1849 (valid
family
of
Oegopsida). Chirothauma Chun, 1910 (valid genusof family CirroteuthidaeKeferstein,1866, Cinoctopoda).
Belosaepia Voltz, 1830; type genus of
ChondrophoraKeferstein,I 866 (invalid grouping).
Belosaepiidae Dixon, 1850,Sepiida).
Cirroteuthoidea(probably meant family Cirroteuthidae
Belosepiella de Alessandri, 1905 (valid genus of Belosepiellidae Naef, 1921,Spirulida). Belosepiinae nov. subfam. (taxon was already proposedby Nyst, 1843 and must be corrected
Keferstein,1866,Cimoctopoda). ClavirostridaeAbel, l9l6 (unavailablefamily name; not basedon a valid genus). Coeloteuthinae nov. subfam. (usually seen as a
to Belosaepiidae,based on Belosaepia Yoltz,
synonym of Passaloteuthididae Naef, 1922; see
1830,Sepiida).
Doyle, Donovan& Dixon, 1994).
BeloteuthidaeWiltshire, 1869 (forgotten name, resurrectedby Riegraf, Janssen& SchmidtRiegraf, 1998, not used by most authors,junior synonymsare Palaeololiginidae Naef, 1921 and Teudopseidae Naef, I 92 1, Vampyromorpha). BelotetrthisMiinster, 1843 (unior, subjectivesynonym
Coeloteuthis Lissajous,1906 (valid genus of Passaloteuthididae Naef, 1922,Belemnoidea). ConirostridaeAbel, 1916 (unavailablefamily name; not basedon a valid genus). ConobelusStolley, 1919 (unior, subjectivesynonym of
R h o p a l o t e u t h i s L i s s a j o u s ,1 9 1 5 , f a m i l y
205 DuvaliidaePavlow.1914.Belemnoidea).
Loligosepiidae Van Regteren Altena, 1949,
Conoteuthis d'orbigny, 11t42(also Conoteuthis Naef. 1922,p.257 : norr. err. pro ConobelusStolley.
vamyroprnorpha).
1926,
Duvalia Bayle. 1878(valid genusof family Duvaliidae p a v l o w ,1 9 1 4 B , elemnoidea).
(Prosch,1847(validfamily of oegopsida). Cranchiidae
puvaliinae nov. subfam. (family name was already proposedby pavlow, 1914).
CrassicarinatiQuenstedt, 1849 (unavailable family name;notbasedonavalidgenus).
E l e d o n eL e a c h , 1 8 1 7 ( v a l i d g e n u s o f O c t o p o d i d a e
1919, family
Diplobelidae Naei
Belemnoidea).
CtenoglossaNaef, 1921 (invalid taxon of unclear hierarchy, about superfamily Bolitaenoidea Chun, 1911; superfamilymust be narled of the oldest available family name which is t i ,L { . r^ u d s dA 'rrnpn 'hri +Lr ra E Hoyle, 1886; therefore
d'orbigny. 1g40,octopoda). E n d o c e r a t i d a eH y a t t , 1 8 8 3 ( v a l i d f a m i l y o f t h e Nautiloidea). Enoploteutlii.s d'Orbigny, 1839 (valid genus of E n o p l o t e u t h i dP ae f e f f e r ,1 9 0 0 O , egopsida).
AmphitretoideaHoyle, 1886. Octopoda). Cylindroteuthinaenov. subfam. (family name was alreadyproposedby Stolley, 1919.valid farnily ofBelemnoidea).
Gastrobelusnov. gen. (valid genus of
family
Passaloteuthididae Naef, 1922,Belemnoidea). GeoteuthisMtinster, 1843 fiunior, subjectivesynonym
C y l i n d r o t e u t l r i sB a y l e , 1 8 7 8 ( v a l i d g e n u so f f a m i l y Cylindroteuthididae
Stolley,
19 I 9,
Belemnoidea).
of Loligosepia
Quenstedt, 1839, family L o l i g o s e p i i d aVea n R e g t e r eA n l t e n a ,1 9 4 9 ) . G. zitteliFraas,1882(junior.subjectivesynonymof Lo I igosepia auIensis (Zieten,I 832)).
Dact.vlotettthis Bayle, 1878 (valid genus of family Megateuthididae Sachs & Naljaeva. 1961, Belemnoidea)
Glt,phiteuthrs Reuss. 1854 (valid genus of family Trachvter.rthididae Naef^1921.Vampromorpha). Conatus Grai'. lttzlg (type genusof GonatidaeHoyle,
DecapodaLeach, 1818(non DecapodaLatreille, 1805, C r u s t a c e ar;e p l a c e db y D e c a b r a c h i B a oettcher.
1886.Oegopsida). G o n i o t e u t h i . sB a y l e , 1 8 7 8 ( v a l i d g e n u s o f f a m i l y
1940, an older synonym of Decabrachia
B e l e m n i t e l l i d a eP a v l o w i n S c h w e t z o f f ,l g l 3 .
Boettcher,1940 is Decembrachiatawinckworth.
Belemnoidea).
1932). DibranchiataOwen, 1832 (most authorsassumethat Dibranchiatais a synonym of coleoidea Bather. 1888, other authors have restricted the terrn to recentforms: Neocoleoidea). Diccelites Bdhm, 1906 (valid genus of Dicoelitidae Sachs& Naljaeva,1967,Belemnoidea). Gustomesow,
19i B,
Belemnoidea). Diploconus Zittel, 1868 (junior. primary homonymof Diploconus
Haeckel. 1860; replacedby
Diplobeltts Naei 1926.lamily Diplobelidae Naei 1926.Belemnoidea). Doratetrthis
basedon a valid genus). Ha.stitesMayer, 1883 (valid genusof tamily Hastitidae Naef, 1922,Belemnoidea). Hastitinaenov.subfam.(validfamilyof Belemnoidea). Heliceras nom. err. pro Helicertis Dana, 1848 (fossil
Dict,-oconites Mojsisovics,1902(valid genusof family Dictyoconitidae
HastatidaeStolley,1919(unavailablefamily name;not
Woodward, 1883 (valid genus of
Plesioteuthididae Naef. 1921.Vampyromorpha). Doratopsis Rochebrune. 1884 (junior. subjective s y n o n y m o f C h i r o t e u t h i sd ' O r b i g n y . 1 8 4 1 , family Chiroteuthidae Gray. 1849.Oegopsida). Dor..vanthe.sMtinster. 1846 (i.alid genus of
fish; fide Riegraf, Janssen& Schmidt-Riegraf, l99g). Aentisepius Steenstrup. 1875 (subgenusof Sepla Linnd, 1758. family Sepiidae Leach, lgl7, Sepiida). H e t e r o g l o s s aN a e f , 1 9 2I ( i n v a l i d t a x o n o f u n c l e a r hierarchy). Hiholites nom. err. pro Hibolithes de Montfort. 1808 (valid
taxon of
family
Mesohibolitidae
Nerodenko,1983,Belemnoidea). H o m a l o t e u t i l s S t o l l e y ,1 9 1 9 ( v a l i d r a x o n o f f a m i l y Megateuthididae Sachs & Nalnjaeva. 1967, Belemnoidea).
206 Idiosepius Steenstrup,1881(type genusof Idiosepiidae
1983and KelaeninaStarobogatov,1983). MetasepiaHoyle, 1885(valid genusof SepiidaeLeach,
Appell
1 8 1 7 ,S e p i i d a ) . Metateuthoidea Naef, 1921 (invalid taxon).
Steenstrup,I 857, Oegopsida).
Myopsida d'Orbigny, 1845 (valid taxon of Kelaeno Mi.inster,1842 (unior, primary homonym of Kelaeno
d ' O r b i g n y , 1 8 4 1 ; r e p l a c e db y
Muensterella Schevill. 1950).
NannobelusPavlow, l9l3
KelaenidaeNaef, 1921 (invalid family taxon, basedon junior,
Teuthoidea).
primary homonym; replaced by
MuensterellidaeRoeer. 1952).
(valid taxon of family
Passaloteuthididae Naef, 1922,Belemnoidea). IrtreohibolitesStolley, 1911 (valid genus of family Mesohibolitidae
Nerodenko,
1983,
Belemnoidea). Leachia Le Sueur, 1821 (valid genusof Cranchiidae OctopodaLeach, 1818(now restrictedto the "incirrate"
Prosch,1847,Oegopsida). Leptoteuthis v. Mayer, 1834 (the original and correct spellingis Leptotheuthisv. Meyer, 1834,type g e n u s o f L e p t o t h e u t h i d i d a e N a e f , 1 9 2 1,
OctopodaLeach,1818). (incorr. spelling of Octopoteuthidae Octopodoteuthidae
Vampyromorpha). Lioglossa(invalid taxon of unclearhierarchy). Lioteuthis
octopods). Octopodidae d'Orbigny, 1840 (valid family of
Berry, 1912,Oegopsida).
Naef, 1922 (1alid genus of family
Lioteuthididae Naef, 1922, Y ampyromorpha). Listroteuthls Naef, 1922 (vahd genus of family MuensterellidaeRoger, 1952,Y ampyromorpha). Loliginites Quenstedt,1849 (unavailable;see Art. 1b and Art.20, "Code"; Doyle proposed
Octopus Cuvier, 1798 (valid genus of family Octopodidaed'Orbigny, I 840, Octopoda). Oc1;thod Rafinesque, 1814 (valid genus of family OcythoidaeGray, 1849,Octopoda). Odontobelus Naef, 1922 (.invalid genus, junior. subjectivesynonymof Acrocoeliles Lissajous.
Jeletzlg,teuthlsas a nomen substitum,which is,
1915, family
however, a new scientific name, since an
Nalnjaeva,I 967, Belemnoidea).
unavailable name cannot be replaced by a nomen substitutum).
Megateuthididae Sachs &
Oegopsida d'Orbigny, 1845 (valid taxon of Teuthoidea).
Loligo Schneider,1784 (type genus of Loliginidae Lesueur,I 821, Myopsida).
Ommastrephes d'Orbigny, 1835 (valid genus Ommastrephidae Steenstrup,1857, Oegopsida).
Loligo sagittata (misspelling for Loligo subsagittata
Ommatostrephes
Agassiz, 1846 (unjustified
Miinster, 1839,junior, subjectivesynonym of
emendationof Ommastrephesd'Orbigny, 1835
Plesioteuthisprisca (Rrippell, 1829), family
and thereforea junior, objective synonym).
Plesioteuthididae Naef, | 92 7, Y ampyromorpha).
Ommatostrephidae(based on the junior, objective
Loligosepia Quenstedt,1839 (valid taxon of the family Loligosepiidae Van Regteren Altena, 1949, Vampyromorpha).
correct spelling is OmmastrephidaeSteenstrup, 1857,Oegopsida).
Megateuthis Bayle, 1878 (valid taxon of the family Megateuthididae Sachs & Nalnjaeva, 1.967, Belemnoidea).
O n y c h i t e s Q u e n s t e d t ,Q u e n s t e d t , 1 8 5 6 ( v a l i d parataxon,basedon isolatedbelemnoidhooks). O n y c h o t e u t f t l s L i c h t e n s t e i n ,1 8 1 8 ( t y p e g e n u s o f Onychoteuthidae Gray, 1849,Oegopsida).
Mesohibolites Stolley, 1919 (valid taxon of family Mesohibolitidae
synonym Ommatostrephes Agassiz, 1846,
Nerodenko,
1983,
Belemnoidea). MesoteuthoideaNaef, 1921 (invalid taxon, taxa which were assignedto Mesoteuthoideaby Naef are now attributed to Teudopseina Starobogatov,
Onychoteuthisprisca Miinster, 1830 (nomen nudum, neverformally published). Orthoceren (polyphyletic grouping of nautiloid cephalopods). Oxyteuthis Stolley, 1911 (valid genus of family OxyteuthididaeStolley, 1919,Belemnoidea).
207 P a c ' h y t e u t h i sB a y l e . 1 8 7 8 ( v a l i d g e n l r so f f a m i l y Cylindroteuthididae Stolley,
1919,
Belemnoidea).
Prototeuthoidea Nael, l92l (usedin its correctedform Prototeuthidaas an order or suborder ol the Vampyromorpha).
Palaeoctopoda Naef, 1921(proposedas a sr.rborder: not usedby Doyle.Donovan& Nixon, 1994). Palaeoctopu.iWoodvn'ard. 1896 (this is a replacement
P s e u c l o b e l u sB l a i n v i l l e . 1 8 2 7 ( v a l i d g e n u s o f t h e family DuvaliidaePavlou,.1914,Belemnoidea). Psetrdoduvalia nov. gen. (valid genus of family
name for Calai'.sWoodward, 1896 which was
DuvaliidaePavlorv,1914.Belernnoidea).
preoccupiedby Calai'sRafinesque.1815:valid
Pseuclohastilesnov. gen. (valid taxon of the tamily
genus of Palaeoctopodidae Do11o. 1912, Cinoctopoda).
Subhastitidae Gustornesor,, 1977.Belemnoidea). PtiloteuthisGabb. 1869 (identitiedas an insectwing.
PalaeololiginidaeNaef, 192I (r,alid genus of larnily
Rehn,1939).
Palaeololiginidae Nael 192l, Vampyrornorpha). P a l a e o s e p i a T h e o d o r i . 1 8 4 4 ( j u n i o r . s u b j e c t i rc s y n o n y m o f L o l i g o . s e p l aQ u e n s t e d t ,i u 3 9 ,
R h u b tlo b e lrl.r nov. gen. (valid genus of family HastitidaeNaef. 1922.Beleurnoidea).
family Loligosepiidae Van Regteren Altena.
R h u p h i h e l u . r l . r o \ ' ._ s e n(. r , a l i d g e n u s o f t h e t a m i l y
I 949, Vampyrornorpha).
Mesohibilitidae Nelodenk.1983.tselemnoidea). R h o p a l o b e l u sP a r , l o u ' .1 9 1 3 ( i n v a l i d t a x o n ,j u n i o r ,
Parabelopehlsnov. gen. (unior. subjectivesynonvlx of Loligosepia
Quenstedt. 1839, family L o l i g o s e p i i d a eV a n R e g t e r e nA l t e n a . 1 9 4 9 .
subiectivesynonylnof Httstites Mayer-Eymar,
Vampyrornorpha).
Belemnoidea).
P a r a h i b o l i t e s S t o l l e y , t 9 l 9 ( v a l i d g e n u so f t a m i l l , Mesohibolitidae
Nerodenko.
19U3.
Belernnoidea). Paraplesioteulfti.sNacf-.192I (valid genusol family
1883, farnily Hastitidae Naef,
1922,
R h o p ( l l o t e u t l l s L i s s a j o u s .i 9 l 5 ( v a l i d g e n u so f t h e t-au-rilv DuvaliidaeParlou', l9 1,1^ Belemnoidea). R o r t d e l c t i o l uN a e t . l c ) 2 1( r a l i d g e n u so l S e p i o l i d a e L c a c h I. 8 1 7 .S e p i o l i d a ) .
Plesioteuthididae Naef'.192I . Vampyromorpha). Passaloteuthinac nov. subfam. (valid family oi the Belernnoidea). P a . s . s a l o t e u t f tLi "i sr s a j o u s ,l 9 l 5 ( r , a l i d g e n u so f t h e family
P a s s a l o t e u t h i d i d a e r . r . - a e l .1 9 2 2 ,
Belemnoidea). P h r a g m o t e u t i l . rM o j s i s i v i s .1 8 8 2( v a l i d g e n u so f t h e f-amilyPhragrnoteuthididae Mojsisovics. I 882. Belemnoidea). Pht'lloteuthi.r Meek & Hayden.1860(invalidtaxon.the type speciesis a plantremain). P l e s i o t e u t h i d a eN a c f . 1 9 2 1 ( r ' a l i c l t a r n i l v o f Vampyrornorpha). Plesioteuthi.\Wagner.1859(r'alid genusof the farnily Plesioteuthididae Naef. I 92 1. Varnpyromorpl.ra) (i nvalidtaxon). Polypodoidea P o l y t e u t h i d aSet o l l e y .l 9 l 9 ( u n a v a i l a b ltca n i i l . vn a m e . not basedon a valid genus). Protodecaptr.l Naef. 1922(unavailablenan.re. basedon a hypothetical animal). ProtosepioitlesNaef. 1922 (unar..ailable narne.based on a hypotheticalanimal). Prototeuthi.s Naef-.i922 (unavailable name.basedon a hypothetical animal).
S u l l t i r t g o t e t t r hLl i s s a j o u s .l 9 l 5 ( v a l i d g e n u s o f t h e t a m i l v S a l p i n g o t e u t h i d i d a eD o y l e . 1 9 9 2 . Belemnoidea). Se1rlaLinnd, 17-58(valid genusof the family Sepiidae L e a c h .1 8 1 7 S . epiida). Sepia tenusta Miinster. 1837 (nomennuduur;species \\'asllcverfbrmallypublished). S e p i u l i t e s M i i n s t e r . 1 8 4 3 ( i n i ' a l i d g e n u s :p r o b a b l y j r . r n i o r .s u b j e c t i v es - y n o n y mo f L o l i g u , s e p i , Q u e n s t e d 1t .8 3 9 ) . SepiidaeLeach. 1817(valid family of the Sepiida,br.ri not authorKef-erstein. 1866). Scpiinae L e a c h .l 8 l 7 ( n o tN a e f. I 9 2 1 ) . S e p i o i d eZai t t e l .1 8 9 5( n o tb y N a e f .1 9 1 6 ) . S e p i o l a S c h n e i d e r l.7 l t 4 ( v a l i d g e n u so f S e p i o l i d a e r L e a c h .i 8 1 7 . S e p i o l i d a ) . SepiertuNaef-,1912(i,alid genusof SepiolidaeLeach 1 8 1 7S . epiolida). Sepiolidae Leach,1817(va1idfarnilyof Sepiolida). Sepiophora Zittel, 1884(invalidtaxon). Sepioteuthis Blainville. 1824 (va1id genus of LoliginidaeLesueur,I 82l, Myopsida). Spirula Lamarck. 1799 (r,alid genus of the family
208 Spirulidaed'Orbigny, I 826, Spirulida). Spirulidaed'Orbigny, 1826 (valid family of the Spirulida).
AcrurotetrthisBerry, 1920).
Spirtrlirostra d'Orbigny, 1842 (valid genus of the family SpirulirostridaeNaef, 1921, Spirulida). Spirttlirostrella nov. gen. (valid genus of the family Belemnoseidae Wiltshire, 1869, Spirulida). Spirtrlirostridae Naef, 1921 (valid family of the
Teuthoidea(valid order,comectedform Teuthida). Teuthopsis Geinitz, 1846 (unjustified emendationof Teudopsis Eudes-Deslongchamps,1835 and therefore a junior, objective synonym, family Teudopseidae Naef, 192l, Vampyromorpha). Thysanotetrllzis Troschel, 1857 (type genus of
Spirulida). Spirtrlirostridium nov. gen. (valid genusof the family SpirulirostridaeNaef, I 92 1, Spirulida).
Thysanoteuthididae Keferstein,
1866,
Oegopsida).
Spirulirostrina rov. gen. (valid genus of the family SpirulirostridaeNaef, I 92 I, Spirulida).
TrachyteuthidaeNaef, l92l
(valid family of the
Vampyromorpha; usually in its corrected form
Spirtiisepia Naef, 1922 (unavailablename,basedon a hypotheticalanimal). Sthenoteuthis
Teuthis Schneider,1784 (: Acroteuthis Berry, 1913 nomennovum (nonAcroteulrls Stolley,1911):
Trachyteuthididae Naef, 192l). Trachytetrthls v. Meyer, 1846 (valid genus of the
Verrill, 1880 (valid genus of
Ommastrephidae Steenstrup,1857,Oegopsida). StylotetrthisFritsch, 1910 (valid genusof the family L o l i g o s e p i i d a eV a n R e g t e r e nA l t e n a , 1 9 4 9 ,
family
Trachyteuthididae Naef,
1921.,
Vampyromorpha). Tremoctopus Chiaje, 1840 (valid genus of the family Octopodidaed'Orbigny, I 840, Octopoda). Tttsotettthis Logan, 1898 (valid genus of the
Vampyromorpha). St1;racotettthls Crick, 1905 (might be the rest of an
Vampyromorpha).
Octocorallia, fide Riegraf, Janssen& SchmidtRiegraf,1998).
VampyroteuthidaeThiele in Chun, 1915 (valid family of Vampyromorpha).
TenuicarinatiQuenstedt,1849 (invalid name,not based on a valid genus! about equivalent to LoligosepiidaeVan RegterenAltena, 1949).
Vasseuria Munier-Chalmas, 1880 (valid genus of the family VasseuriidaeNaef, 1921,Sepiida). VasseuriidaeNaef, 1921(valid family of the Sepiida).
TettdopsisEudes-Deslongchamps, 1835 (valid genusof the
family
Teudopseidae Naef,
1921,
Vampyromorpha). Teuthidae(invalid taxon,formerly usedfor Teuthida).
XiphotutnhisHuxley, 1864 (invalid genus, junior, subjectivesynonym of Atractites Grimbel, 1861, family XiphoteuthididaeBather in Blake, 1892, Belemnoidea).
209 List of new species:
References:
A c ' a n t h o l e t t t h i .. j;u e t ' k e l i 1 1 o v .s p . ( " P h r c r g n t o t e t t lh i s "
D o y l e .P . ; D o n o v a n D . . T . ;N i x o n . M . 1 9 9 4 .P h y l o g c n y
monteJiore i ( Buckrnan. I 880). Acanthoteuthi,s problematicd
and svstematics of the Cclleoidca. The Universityof nov. sp. (indet.
belemnoid).
KansasPalcontologicalContributions,ner,vseries. 5 : l - 1 - 5L. a r . v r c n cKea. n s a s .
B eI emno s i s r:of?tnta n n i no\'. sp. (B eI entn rt :;i,; L()s,\tttLtt tt t i Naef, 1922). Belemnotettlhi.s uttrtu no\'. sp. (Chortdt'otetrthi,s r.tturtr (Nael-.1922)).
R i e g r a f ,W . ; J a n s s e nN. . ; S c h m i d t - R i e g r aCl ' . 1 9 9 8 . C e p h a l o p o dd ai b r a n c h i a t a l b s s i l e s( C o l e o i d c a I)I . Fossilium C a t a l o g uAs n i m a l i a .l 3 - 5 5 : - 51 2 ,L e i d c n .
Beloptera longct nov. sp. (Beloptet'a /origa Nacf. 1922). Belopteridiutn ptterilis no\'. sp. (Beloptat'iditrn puerili.s Nacf" 1922). C eI aenotetrt h i.si n c'ert o no\'. sp. (.Cc I u en tt t etrt h i,s i ttc'er I u Nacf. 1922). Lioteulhi.s
C L r t t l c f i s h eOsc. t o p u s c sa.n dA l l i e s . l - 3 5 1 . T r o p i c a l F i s h H o b b y i s t P u b l i c a t i o n s I. n c . . N e p t u n e C i t y , Nerr Jcls,r,.
problentuti('./ nov. sp. (llo teurhi.t
pro b I e rttut it tLNae1" I 922). Paraples ioleuth i.\ nldgn(/ no\'. sp. (Pat'crplesiotattthis sugitteta (Mtinster. 18.13). Spirulirostrct
N e s i s .K . N . 1 9 8 7 .C e p h a l o p o dosf t h e W o r l d : S q u i d s .
.sepioitleu nov. sp. (Spiru I it'u.';rtt
.sepi o i dea Naef . 1922 ). Sp iru I i ro s tt'i cliu tn obtu s unt l1o\.. sp. ( Sp i ru I i ro.t I r'i cli trnt o b h r s u r tN t a e f" 1 9 2 2 ) .
S u ' e e u e vN. { . J . ;R o p e r .C . F . E . 1 9 9 8 .C l a s s i f i c a t r o n , t l p c l o c a l i t i e s "a n d t v p c r e p o s i t o r i e so f R c c c n t C c p h a l o p o d aI n. V o s sN A " \ ' ' e c c h i o nM e . Toll RB, ( e d s ) S n c c n c r '\ ' 1 . 1 S v s t c n t a t i casn d b i o g e o g r a p h y o l - c c p h a l o l . r o cSl sn.. r i t h sC. o n t r .Z o o l . 5 8 6 ( 2 ) :5 6 1 -599
