Die fossilen Tintenfische. Eine paläozoologische Monographie. Fossil dibranchiate cephalopods. Band 5 [PDF]

Zitiervorschau

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 i : i :

;, _o ^

!

I&\

t

ffi

:1 t

t#gt

ffi 'ffi

-

-

> !

'

:

-

=

i

\

.

-

J

c . 3 = ; E / '

=

2

>

=

i

:

: z i? i E ;

i

+

j ? . J t =Z_ i= :? Z, ; EE - . =

ita_l

= ; 2 F

z ,

7 - : _

-

-

=

9

:

-

=

?

=

1a,a?= i

f+rj f

=

:

tffi

ii .l

I

T

l: = = i l - : E= = a 2|

iii\

'j

;

E

;' ' J, . 7i . 4i iZ l= 1 F --

til

4tl

E Z i

'! " . =

3 E : = E E V 7 : : i = t : : = } . E

' = r , ' 7 . = - - ; .

1

v

.

-

.

a t a ? z ; = i - - - !

1 =

11

=

0

? =

'

t

i

0

=

= = i ' - 2 7+ E a : = = - 7 a t t !

'

=

- ? =

.

i

3 . z r i = : 1 ? =

/a

)y r= t =r := z. z_ ri ,. zt ! o

1t l 1 It{1

is)

-

=

.

=

-

=

L

-

-

2

t

=

.. = _,-: 7 = = i - Z

=

r

.

=

?

:

a

F

-

i

;

-

=

-

r

t

=

?

:

= t Z

=

c

:

i

i

t ?: E

z \ t - Z : > = -

=

'

>; = . .

i r ; . ' = - a = ! ; i . - = - ? t s ! 2 i = 2 1 /

\:/

c

7

-

-

2 = 7 , . - . ' - A . ? - = = = 4 z Z y a = i l r . ? l Y Z z = E = = , ;Z, t r = 7 ! - . t r = l : - = = ' - = -

2 = '=" iv -, .l 2: F= i

= : ; 2 4

; a t : 4 : = - a - d

== >= E

-' -: !; € 3 5 . 3 =

" a a : . 2 4 5 2 , 2 t = 2 . = = ; + + : = 4 :- 2 , ' E t

, t

E E = e

7=

, -

= =

= : 2 i . 7 = i ;g =t ; :

= : . =

E 2 > . i - i = l = E 3 i ' , - = = = =

:

= - . - - i L l r . ' z ?

_

!

=

=

^

=

-

=

-

t

,

=

=

-

7

-

,

.

3

_

= J = ; = = ? p

-

:

1

f

.

j

,

=

:

=

=

'

,

i = y = . _ . e ; i e = E ; at E i 2ze _ > = ; ; 3 ; " . ! + ; , L E z E = " E 5 : : >

.

2

'

L

=

u

r

=

.

2

i

=

?

.

=

=

= t = F , a ; t - = = ? i ; u - - r , 6 = a = : =

-

.

a

=

F

/

-

s

=

.

!

2

=

.

J

=

-

-

-

-

. ;

. = E = E E a = t 7? = 1 = "

-

;

t

u

6

=

+ I ; E : E ' = i E 1:

S i - : s i z a = E € : . 5 |? ' : = y : : V y , - t l 7 r = ! ' =

^ r a > Z Z _ . E : . i Z ! - -

: i : 7 = r ; i i ; , = =aa

6 cn FI F

@)

1 , 3Z L ' :i Z= + L 9 a Z i ; . , -: 9; i It : 9 != ; = + € : t ' - E Z : . i E + -

b

ij

A a E = T =e , i 1 i , Z = . = , i

: i j a L T z E i i E E * E ,

= ; a E z : ? v 2 j e , ' j i j

z' L' ; i . =' i; "/ := ; == = 1 ; : j = E t * i i : = . . . - = ? 3 = = i , 5 = Z a ; = a = 1 . = u

-

.

i

'

-

;

u

!

.

c

c

)

=

'

=

=

=

-

i

i

=

= ' , t a l r j = 3 : V i y i ; i . 1 : =

? . : Z = 5 ' - u i

n i

=

=

-

/.

-

-

:-:

j-:

;

=

;:

g

.

=

3 : = 4 = . " e ! 3 = : = l + . * : t r z z ' = t 3 1 a i F 3 : 6 j . T i = + l l = : 7 q . t a 1 - . 2 t

r

c

J

i

;

=

7

:

i

\

!

-

-

=

r

-

102

,i

tl"l

;tb

,:|n4

: h

{. J \\

',tril",

'ii'x',{r,,,

I

rIr

g

",{i\]i

k -t€

,r\t l t\ i.

\:

i:

. - - i.\\\ \ { l\ . '

is\..

\\ \\

|i

l'

ii

i

I

I

Y

l

I I

-

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

\\ 'A4

l,{ t1

l

t.4

Il ' 4i l 1 l

l 1 l

\\7/

i{

t{

lfi / a

\VI

IAt i \l l 1 ul t l { \ d1 lt 1.{ t{ t f \,"1 V Y V , o1" ( e @ ail,6 ;@;@;@; a) \/

\

t

l

\

v

'\s{r

t

\

{

l

4

\t

{ o i

oo

{

o ( i o

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')

)$JJ

/P

\/\

vI I

b

/r

\

r

"

%,, i

\'lu

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

3s€

qt rl' !' -g

E EE o o = 3:€

a

c ) O E

OcD
: O :

5s &

:: E : ; 9

:ot .:g

o

o

(,

!

-r€o €o)

qt fo

f

..)

o (o

g3EgEi tl

o a!

(D t!

=

o)

c u t

(t

o 6

O F

E

s € E€

l f o * : =

.c gg t'l

o CI

-

.=

t

rg

ct o . p

{ ! E 1 5 5

A to I

o, E S e .: X(/ro, J J O . . ( q

a) ( L

.r, a

ot (o

:g f c) o

!

p L q

L

C t

o (o g L

qt

5 L

o

.= t ': 'ct n

ta L

:Gl

(ol

!

t.'|l (nl

(t

#

r

ltc t.h 1.4 I O l o

cr--.-..-_ (o 'trtl o+ q (1)

p

Cen.

o Gauli, }C #

c Neoc. 3 7 a

d

i O l roF l* (Jt-*

-oo\\\\ oa)ll (oi oi \\ € t\\ ( o l o2 \ C _ l e\ -o-a

6

- ll f r \ \ \ e\ig \\,,j 'TE- , -\I

I

e i $ ; i ?qE /F €,c

EE 2,., rF l6--E F

d

-

q

e

c

d

v p d

Rhat. Non, Karn. Lad. Anis. Skyt.

-

E--J t l

(

eiTT sT'l

El, e\ I s:t t:u ll*

//.--

l l o

//s

z \\ ,tE

eo \\

E

v p

o

t' t t d

e o

o

ii"A

a

d

o

X x o \ \

v p o o o o

CD

' 'Ttl :$l- t ro to a

g- oa? \\\ \ l s

\\

|

+-t """H+ "r\r

\

\

A o

{#

=

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.

S h e l l s .C e p h a l o p o d aR. e p . B r i t . A s s . f . a d v . o f

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

Sc.(cf. 1844,1847.)

Anirnal.Qu. J. Geol.Soc.,v. LXXIII. 1 8 8 8 - 9 1 .C r i c k & F o o r d , C a t a l o g u eo f t h e f o s s i l

1914.Chapman.F.AustralasianFossils.London. 1859. Chenr.r,J.C., Manr,relde Conchyliologieet de Pal6ontologie Paris,Masson. conchyliologique. 1 9 0 0 . C h u n ,C . . A u s d e nT i e f e nd e sW e l t m e e r e sJ .e n a . ( 2 n de d i t i o n 1 , 903.) 1910. Ibid., Spirula australi.sLam.

Cephalopodain the Brit. Mus. (Dibranchiata, Tetrabranchiata). 1893.Darnes, W.,

Uber

die

Gliederung der

f lozformationenHelgolands. Sitz.-Ber. K. Ber. math.-phys.

Kl.Siichs.Ges.Wiss.,Vol.LXII. 1910. Ibid., Die Cephalopoden.I. Teil: Oegopsida. W i s s .E r g .D . T i e f s e e - E x pV. ,o l . X v l l l . 1915. Ibid., 2. Teil: Myopsida,Octopoda.Vol. XVIII 2. 1 8 9 4 . C l a r k e ,J . M . ,T h e e a r l ys t a g e so f B a c t r i t e sA. n n . Geol..v. XIV. 1 8 9 5 - 9 8C . o B m a n nM , . , M o l l u s q u e se o c 6 n i q u edse l a Loire- Inf-erieure. Boll. Soc. Sc. de I'ouestde la Fr. t. 5. Nantes. 1900-03.CoBmann,M. & Pissaro,G. Faunedocenique d u C o n t e n t i n ( M o l l u s q u e s )B u l l . S o c . G 6 o 1 . N o r m a n d i et ., 1 9 e t 2 3 . 1910-13.lbid., lconographiecornpldtede coquillesde I'eocdnede environsde Paris.t. 2. Paris. 1896. Crick, G.C., On the proostracumof a Belernnite

preuB.Ak. Wiss.Berlin,phys.-math. Kl. 1 9 0 1 . D e a n , B . , N o t e s o n l i v i n g N a u t i l u . s .A r n e r i c . Naturalist,v. XXXV. 1835.Deslongchamps, E.,

Mdmoires sur les

Teutopsides.Mem. Soc. Linn. Normandie,t. 5. 8u11. Geol. 1824. Deshayes,G.P., Decription des Coquilles fossilesdesenvironsde Paris.t. IL Ibid. 1837. 1 8 3 1 . I b i d . , C o q u i l l e s c a r a c t e r i s t i q u edse s t e r r a i n s . Paris. 1839-57.Ibid., Trait6 eldrnentaire de Conchyliologie.t. III. Paris. 1 9 1 5 . D i e n e r , C . , U b e r d i e B e z i e h u n g e nz w i s c h e n A u l a c o c e r a . sH a u e r ,A s t e r o c o n i t e sT e l l . u n d DictyoconitesMojs. Sitz.-Ber.Ak. Wiss. Wien, Vol. CXXVI. 1912. Dollo, L., Les Cdphalopodesadaptds2rla vie

189 n e c t i q u e s e c o n d a i r ee t d 1 a v i e b e n t h i q u e tertiaire.Z. Jahrb.Suppi.15,Vol. l. 1891. Duval-Jouve, J., Belemnitesdesterrainscrdtacds

t2. 1 8 5 3F . rischmann,

L.,

Versuch

einer

Z u s a m m e n s t e l l u n gd e r b i s j e t z t b e k a n n t e n

infdrieursdes environsdes Castellane(Basses-

fossilen Tier- und Pflanzeniiberreste des lithogr.

Alpes)

Kalschief-ersin Bayern. Progr. Eichstatt.(Altere

concid6rds gdologiquement e

z o o l o g i q u e m e n ta, v e c 1 a d e s c r i p t i o nd e c e s t e r r a i n sC. . R .P a r i s . 1849. Edwards,F.E., Monographof EoceneMollusca. Part.L Cephalopoda. Palaeontogr.Soc..London. 1 8 7 7 . E d w a r d s ,F . E . & W o o d , S . V . , M o n o g r a p ho f

Literatur!) 1 8 7 2 . F r i t s c h , A . , C e p h a l o p o d e nd e r b c i h m i s c h e n Kreideformation. (Unter Mitwirkung von H. Schldnbach.) 1910. Ibid.,Miscellanea paleontologica. lI. Prag.

EoceneCephalopodaand Uivalvesof England,

1861. Gabb,W.M., .!r.,nopsls Moll. Cret. Form. 32.

Pal. Soc.London.

1 8 6 4 - 6 9 . G a b b ,W . M . & M e e k , P a l a e o n t o l o g yo f

1124. Ehrhardt.B., De belemnitisSuevicisdissertatio, qua in primis in obscurihactenusfosilis naturam

Califomia.L u. II. Geol.Surv.Calif-.

inquiritur. Leyden. (First to recognizethe typical

1868. Gastaldi,B., (tber SepicrMichelottii i) Mem. A c c a d .S c .T o r i n o( 2 ) t . 2 4 .

similaritywith Nautilus,hencewith the general

1904.Gemmellaro,G.S.. I Cefalopodi del Trias

cephalopodfeaturesl) 1 8 9 6 . E n g e l . T h . . G e o g n o s t i s c h eW r e g w e i s e rd u r c h Wtirttemberg.Stuttgart. 1906. Felix, Joh.,Die Leitfossilienaus dem Pflanzenund Tierreich in systematischerAnordnung. Leipzig. 1 8 3 5 . F d r u s s a cA, . E . . E r s t e rT e i l d e r , , C 6 p h a l o p o d e s

superioredella regioneoccidentaledella Sicilia. Palermo. 1 8 4 7 - 5 2 .G e i b e l . C . S . , F a u n a d e r V o r w e l t . L e i p z i g (Literatur!). 1 8 5 2 . I b i d . , D e u t s c h l a n d sP e t r e f a k t e n ( L i s t e r n i t Synonymaund Fundorlen). 1910. Grandjean,F., Le siphon des Ammoniteset de

ac6tabulifdres". Paris.(F6r. et d'Orbigny,A. de,

Bdlemnites:Bull. Soc.gdol.France(a), t. l0 (cf.

Histoire naturelle gen6raleet particulidredes

C . R . P a r i st,. 1 5 0 ) .

Cdphalopodes ac6tabulifdres vivants er fossiles. P a r i s 1 8 3 5 - 1 8 4 8( g e n e r a l l y 1 8 3 9 a n d b y d'Orbignys!). 1887. Fischer.P.. Manuelde Conchyliologie. Pans. 1882. Fraas, E.. Loliginites (Geoteuthis) Zitelli E. Fraas.Ein vollstzindigerhaltenerDibranchiate aus den Laibsteinendes Lias s. Jahreshefte Ver. vaterl.Naturk. Wr.irlternberg, Vol. XLV. 1 9 1 0 .I b i d . , F i i h r e r d u r c h d i e K g l . N a t u r a l i e n sammlungzu Str.rttgart, 3rd edition,Stuttgart. 1 8 5 0 . F r a a s . O . , V e r s u c h e i n e r V e r g l e i c h u n gd e s deutschenJuras mit dem franzcisischenund englischen.N. Jahrb.1-.Min. 1855. lbid., Loliginite,salatus. Wr.irttemb.Jahreshefte ll. 1866. Ibid.,.,Vorder Stindfluth".Stuttgart. 1 8 7 8 .l b i d . , A u s d e m O r i e n t . I I . T e i l . G e o l o g i s c h e Beobachtungen am Libanon. Stuttgaft. 1 9 0 8 .F r e c h , F . . L a t h e a g e o g n o s t i c a . 2 . D a s Mesozoikum. Vol. I. Trias.Stuttgaft1903-08. 1868. Frirdn,P., Quelquesmots surune Bdlemnitedu Lias n-royen. Metz Bull. Soc.Hist.N. Moselle.v.

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

XVI.

I

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

1889. Ibid., Observationson the anatomy of rare Cephalopod(GonattrsFabricii). Proc. Zoo1. Soc.London.

Kopenhagen L Abh. d. n. Ges.Wiss.,Vol. XXII. Giittingen. 1892. lbid., Das norddeutscheUnter-Oligociin und

1864. Huxley, Th., On the structureof Belemnitidae... Mem. geol. surv.Un. Kingdom. 1886. Huxley, Th. & Pelseneer,P., Report on the specimen of the genus Spirula collected by H.M.S. Callenger. Chall.Rep.,v. XLIV.

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.

Anatomie der wirbellosenTiere. III 1. Mollusca.

Bui. Mus. Comp.Zool. Cambridge, v. III. .l 890. Jaeckel (Uber Ac:anthoteuthis). Sitz.-Ber.Ges.

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

some

Belemnites and other fossil remains of Cephalopoda in

the

Oxford-clay near

Towbridge, Wiltshire. Philos. Trans. (Ibid.

paleocrdtacdes de sud-ouestde la France.Mem.

Supplementaryobservations1850). (cf.: Ann.

pour servir d l'explic. de la carte gdol de la

Mag. N. H. (2), v. X.)

France.

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 . .

Filiation

der

B e l e m n i t e , gc t c u t i .

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

Solnhofen.KefersteinsTeutschland, Vol. V. 3 p.

County.U. S. geo1.Surv.of the territories. v. IX.

5gl.

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

1836. Ibid., Mitteilung an Prof. Bronn. N. Jahrb. f.

Coprolithen in Deutschland.N. Jahrb.f. Min..

Senkenbergianum L Min.

Belemniten.Bayreuth.

Vol.I.

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.

of Newton and Wautubbee,Miss. Journ. Cinc. Soc.Nat' Hist.,v. IX. 1 8 4 7 . M i c h e l o t t i , G . , D e s c r i p t i o nd e s f o s s i l e s d e s terrains miocdnes, de I'Italie septentrionale. Leyden.

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.'

1834, 1836. lbid.. Brielliche Mitteiiunsen an Bronn. N. Jahrb.f. Min.

II. Zool.

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

systdmatiqr.re et

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.

Formen als Urkunde ihrer Stamrnesseschichte.

Reichsanst. Wien, Vol. X.

Jena.G. Fischer.

1 8 8 5 .I b i d . ,

Uber

das

Belemnitengeschlecht

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.

und

1919.Ibid.. Bilder vom Bau und Leben der Tintenf-rsche. Vierteljahrsschr. Ziiricher nat. Ges. 1 9 2 0 . I b i d . , U b e r d a s s o g e n a n n t e, . b i o g e n e t i s c h e Grundgesetz". Festschr. f. Zschokke.Basel. 1 9 2 1 .I b i d . ,

Das

System der

dibranchiaten

Cephalopoden und die mediterranen Arren

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.,

Uber Bau und Entwicklung des

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

rrr,1852).

Palaeontology. London and

Edinburgh.

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

preservedwith a GreatPortion of their Soft Parts

ventouses dans le rdgne animal. Rec. zool.

in the Oxford Clay. Philos.Trans.London. (cf.

Suisse.t. 2.

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

of

Dibranchiate Cephalopoda

(Coccoteuthislatipinnis)from the upper oolitic

Alpes suisses.Cataloguedes Cdphalopodes des

shalesat Kimmeridge.Quart. Journ.geol. Soc.

Alpes suisses. Gendve.

London,v. XI.

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

(2ndedition1861). 1860. Ibid.,Palaeontology

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

Jahreber.ibid.

der Loligeneen oderTeuthidae

1909. Stromer v. Reichenbach, E., Lehrbuch der Paliiontologie. I. Teil: WirbelloseTiere.Leipzig und Berlin.

(Owen)

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

Cenomanbildungen von Podolien... Extr. Bull. Acad.Sc.Cracovie.Classesc.math.nat.

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 ,

A..

Uber

einige

Belemniten

Wrirtternbergs. Jahreshefte Ver. vaterl. Naturk. Wiirrt.Vol.l. 1678. Lister.M.. Historiaanimalum.Londini. 1 8 9 8 . L o r e n t h e y . E . . S e p i a i m r . r n g a rT. e r t i r i r .( S . ltt.rytguria 1. sp.) Math. nat. Ber. Ungarn. Vol. 15.

et 3. (Erweiterung desWerkesvon 18241)

tSZ7. Michelotti. J.. Obser.r,ation sur unc Argonaute fossile.Ann. Sc.Nat. (2) t. g. Zool. paris.

1874. Dumortidr. E." Etudespaleonologiquessur les

1834. Mrilster. G.. Craf zr.r.Uber zrveineueArten von

dep6tsjurassiqr.res dr-rbassinde Rh6ne.v. 2-4.

Sepla.Stuttgart. 1899. Oppenheirn,P., IJber Orcagnia trit,igiartan. g.

Paris1864-74 1 8 3 5 . F d r u s s a cA , .E.de. Lettre sur les Belentnites. a d r e s s d ed M . l e P r c i s .c l e l ' A c a d . R o y d e s

e t s p . .e i n e nn e u e nd i b r a n c h i a t eCne p h a l o p o d e l . Zeitschr.d.D.geol.Ges.,Vol.5l.Verhandl.

t96 1 8 9 2 . P a r o n a , C . F . , D e s c r i z i o n ed i a l c u n i f o s s i l i miocenicide Sardegna.Atti Soc. Ital. Sc. Nat., v. 34. Milano. du Jura et du 1913. Pavlow, A.P., Les Cdphalopodes Cr6tac6inf. De la Siberie septentrionale.M6m. Ac. Imp. Sc. Petersbourg. 1865. Phillipps, J., Note on Xiphoteuthis elongata. G e o l .M a g . ( l ) v . 2 . L o n d o n . 1901. Reis, O.M., Eine Fauna des Wettersteinkalkes,

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.

Soc. Imp.

NaturalistesMoscou.t. 34. 1872. Vincent, G., Un Belemnosepiaet un Cerithium nouveaux pour la Faune Bruxelliennes.Ann Soc.Malac.Belgique.t. 7. Bull. p. CXVII. 1 9 0 1 . I b i d . , C o n t r i b u t i o n s d l a P a l d o n t o l o g i ed e

1 . T . C e p h a l o p o d e nG . e o g n o s t .J a h r e s h e f t e .

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 .

Jahrg. I 3 ( 1900) Miinchen, (cf. Jahrg. l8

Ann. (M6m.) Soc.Mal. Belg.t. 35. Bmxelles.

(1eOs)). 1813. Schlotheim,E.F.v.,Beitr. z. Naturgeschichte der Versteinerungen in geognost.Hinsicht. Leonh. Taschenb. f. ges. Mineraiogie. 7. Jahrg. Frankfurt.

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.

1824. Stahl, Ubersicht iib. d. Versteinerungen Wtirttembergs ... Corr.-Bl. wrirttemb. landw. Ver. Stuttgafi-Ttibingen.

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