SI-IELL ULTRASTRUCTURE,

MUSCLE-SCARS, AND BUCCAL

APPARATUS IN AMMONOIDS

L.A. D O G U Z H A E V A
Paleontological Institute of the USSR Academy of Science, Profsojuznaya 123, Moscow 117868, Russia.
H. M U T V E I
Swedish Museum of Natural History, Department of Palaeozoology, S-10405 Stockholm, Sweden.

DOGUZHAEVA L.A. & MUTVEI H. 1993. Shell ultrastructure, muscle-scars, and buccal apparatus in ammonoids.
[Ultrastructure de la coquille, empreintes musculaires et appareil buccal chez les ammono'~des]. GEOBIOS, M.S.
n" 15 : i i i - i 1 9 .

ABSTRACT
(1) The Cretaceous genus Aconeceras has large ventro-lateral muscle-scars in addition to smaller dorsal and ven-
tral scars. This indicates that some ammonoid groups at least probably had strong retractor muscles to the head
and funnel, and that their muscle system was more complex than that in the Recent Nautilus. (2) Radula of
Aconeceras has multicuspid teeth, known among recent cephalopods only in bathypelagic octopods. (3) In the
Cretaceous heteromorphic genus Ptychoceras, repetition of shell layers and mode of truncation of initial portion of
shell indicate that the shell surface was covered by soft tissue, at least in apertural region. (4) Shape of shell
aperture in many ammonoids differs considerably from that in Nautilus by lack of hyponomic and ocular sinuses.
This may indicate principal morphological differences in the head-funnel region of the body between ammonoids
and Nautilus. (5) Transition from retro- to prochoanitic septal necks, which took place with different evolutionary
rates in different ammonoid orders, improved the osmotic pumping system, and made it possible to form a new
portion of siphuncle prior to secretion of next septum.

KEY-WORDS : MUSCULATURE, SHELL ULTRASTRUCTURE, BUCCAL-APPARATUS AMMONOIDEA.

RC,SUMt~
(1) Le genre cr~tac~ Aconeceras poss~de de longues empreintes musculaires ventro-lat~rales qui s'ajoutent ~ de
petites empreintes lat~rales. Cel~ indique que certains groupes d'ammono~des, au-moins, ont de puissants muscles
r4tracteurs pour la t~te et l'entonnoir et que leur syst~me musculaire ~tait plus complexe que celui des Nautilus
actuels. (2) La radula d'Aconeceras poss~de des dents multicuspides qui ne sont connues que chez les octopodes
actuels bathyp~lagiques. (3) Sur le genre h~t~romorphe cr&tac~ Ptychoceras, la r&p~tition des couches du test et le
mode de troncature de la portion initiale de la coquille indiquent que la surface ~tait couverte par des tissus mous,
au-moins dans la r~gion aperturale. (4) La forme de l'ouverture de nombreux ammonoides dif~re consid~rable-
ment de celle du Nautilus par l'absence des sinus hyponomique et oculaire. Cel~ indique que la morphologie de la
t~te peut presenter des differences essentielles entre les ammonoYdes et le Nautilus. (5) Le passage de la structure
retrochoan~e A la structure prochoan~e apparait avec des taux ~volutifs diff~rents dans divers ordres d'ammono~-
des. Cel~ s'accompagne d'une amelioration du syst~me de pompage et rend possible la formation d'une nouvelle
portion de siphon avant la secretion de la cloison suivante.

MOTS-CL]~S : MUSCULATURE,ULTRASTRUCTURE,APAPREIL BUCCAL, AMMONOIDEA.

 112

INTRODUCTION ammonoid morphology, such as the ultra-struc-

ture of the shell wall, the development of the pro-
The present paper summarizes some of the re- choanitic septal necks, the muscle-scars, and the
sults of our joint studies on the morphology and buccal apparatus (Doguzhaeva & Mutvei 1986a,
structure of ammonoid shells from the USSR. 1986b, 1989, 1990, 1991, 1992).
The ammonoid material at our disposal, collected
from the Middle Triassic (Anisian) of the N. Cau-
casus and the Lower Cretaceous (Aptian) of the MUSCLE-SCARS
Volga River and the N. Caucasus, is remarkably
well preserved. This has made it possible to stu- Dorsal and ventral muscle-sears have been des-
dy several still inadequately known features of cribed in several ammonoids (for references see

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Figure 1 - a - d : shell muscles in A c o n e c e r a s . a,b : outline and extension of the ventro-lateral muscle-scar (vls) at a late and a n
early ontogenic stage, respectively ; c : reconstruction of the hyponome retractor (hr), cephalic retractor (cr), dorsal muscle (din),
ventral muscle (vm), and the cephalic region of the body with a large hyponome (h) and aptychus (a) ; d : a n n u l a r a t t a c h m e n t of
the body (an) to the shell in front of the last septum, projected to one plane ; ventro-lateral (vls), v e n t r a l (vs) and dorsal (ds)
muscle scars black, e : a n n u l a r a t t a c h m e n t of the body (an) to the shell in N a u t i l u s , projected to one plane ; muscle-scars of t h e
cephalic retractors (rm) and the dorsal a t t a c h m e n t site (d) black, a-d : muscles coquilliers d'Aconscerau, a, b :forme et exten-
sion de l'empreinte ventro-latdrale (vls) dans les stades ontogoniques prdcoces et tardifs ; c : reconstruction du rdtracteur de
l'hypon6me (cr), du rdtracteur edphalique (cr), du muscle dorsal (dm), du muscle ventral (vm) et rdgion cdphalique munie d'un
large hypon5me (h) et d'un aptychus (a) ; d : fixation annulaire du corps (an) & la coquille en avant de la derni~re cloison, projetde
en plan ; en noir : empreintes ventro-latdrales (vls), ventrales (vs) et dorsales (de). e : fixation annulaire du corps (an) sur la
coquille de N a u t i l u s en projection plane ; en noir : empreintes des rdtracteurs c~phaliques (rm) et fixation dorsale (d).
 i13

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Figure 2 - Buccal a p p a r a t u s i n A c o n e c e r a s . a : Tangential section of t h e living c h a m b e r (sw) to show t h e position of aptychi (a)
and counter aptychi (ca) ; b : detail of a section of anterior portion of a n aptychial valve to show t h e calcified anterior edge (ac),
t h i n i n n e r lamella (il), i n n e r calcareous layer (cl) a n d outer organic layer (ol), x 450. c : r a d u l a r t e e t h 9 rachidial (R), lateral (L1,
L2), m a r g i n a l (M1) and m a r g i n a l plate (MP1). d : ctenoglossal radula in the incirrate octopod J a p e t e l l a (from Nixon 1988, fig.
5b). Appareil buccal d'Aconeceras, a : section tangentielle de la chambre d'habitation (sw) montrant la position des aptychus (a)
et des contre-apthychus (ca) ; b : ddtail d'une section de la partie antdrieure d'une valve aptychiale, m o n t r a n t le bord antdrieur
ecdcifid (ac), la fine lamelle interne (il), la couehe calcaire interne (il) et la couche organique externe (vl), x 450. c : dent radulaire :
dents radiales (R), latdrales (L1, L2), marginales (M1) et plaque marginale (MP1). d : radula ctenoglosse de l'octopode J a p e t e l l a .

Jordan 1968). An additional type of muscle-scar, at the Volga River, and in the Callovian ammu-
the ventro-lateral scar, was found in the Aptian nite Quenstedtoceras from Lukow, Poland (Do-
ammonites Aconeceras and Deshayesites collected guzhaeva & Mutvei 1991). The ventro-lateral
 114

muscle-scars form a prominent, adaperturally-di-
rected lobe on each side of the living chamber
(vls, Fig. 1 a,b,c,d). This lobe is subdivided into a
ventral and a dorsal minor lobe. The dorsal mi-
nor lobe may have been the area of origin of the
cephalic retractor muscle (cr, Fig. lc) which ex-
tended to the cephalic region, whereas the ven-
tral minor lobe may have been the area of origin
of the hyponome retractor (hr, Fig. lc) muscle
which extended to the hyponome (h). In Recent
Nautilus the hyponome retractors are weakly de-
veloped, but they are much stronger in Recent co-
leoids, and may also have been strongly develo-
ped in ammonites. In Aconeceras also a p~ired
dorsal and unpaired ventral muscle-scars occur
(ds, vs, Fig. ld). The dorsal muscle (dm, Fig. lc)
probably functioned to draw the head into the li-
ving chamber and to attach the body to the shell,
whereas the ventral muscle or ligament (vm, Fig.
lc) seem to have been used to maintain the
shape and position of the circumsiphonal invagi-
nation in which the prochoanitic neck and the
connecting ring was secreted.
In the material at our disposal we have also ob-
served muscles-scars in the following Jurassic
and Cretaceous genera, Phylloceratida : Holco-
phylloceras (ventral), Euphylloceras (ventral) ;
Ammonitida : Virgatites (ventral), Elatmites (ven-
tral), Dorsoplanites (dorsal), Olcostephanus (ven-
tral), Zurcherella (dorsal), Nodosohoplites (ven-
tral), Melchiorites (ventral), Diadochoceras (ven-
tral), Baculites (dorsal) (unpublished observa-
tions).
As in Recent Nautilus, the posterior portion of
the boqly in ammonoids was attached to the shell
wall along a myo-adhesive epithelial zone (annu-
lus) comprizing areas of origin of the body mus-
cles (an, Fig. ld,e). On the shell wall, the at-
tachment site of the annulus is usually charac-
terized by the formation of the inner prismatic
layer, which corresponds to the prismatic mantle
adhesive layer in Nautilus (Doguzhaeva &
Mutvei 1986a ; Landman et al. 1989). Moreover,
also in Nautilus the mantle margin is attached to
the apertural margin of the shell, secreting the
prismatic mantle adhesive layer which invests
most of the living chamber. The latter layer pro-
bably corresponds to the wrinkle layer which oc-
curs in several ammonoids and fossil nautiloids
(Doguzhaeva & Mutvei 1986a). However, in
Aconeceras, and also in the heteromorphic Pty-
choceras, the mantle seem to have extended, at
least in the apertural region, to the outer shell
surface, thereby providing a more intimate at-
tachment to the shell (Doguzhaeva & Mutvei
1989, 1991).
In several genera of Mesozoic Ammonitina, as
well as in Aconeceras, the shell aperture forms a
sharply pointed rostrum and lateral lappets. On
the other hand, in N a u t i l u s and fossil nautiloids
the shell aperture possesses hyponomic and ocu-
lar sinuses. This may indicate that the cephalic
region in many ammonoids was considerably ex-
tended in front of the shell aperture, and t h a t the
hyponome (h, Fig. lc) was large and therefore
needed to be supported by the rostrum (Do-
guzhaeva & Mutvei 1991).
BUCCAL APPARATUS
Preservation of ammonoid radulae requires ex-
ceptional conditions. Therefore, the radula was
hitherto known in some detail only in two gene-
ra: in the Carboniferous goniatite Eoasianites
(Closs 1967) and Jurassic ammonite Elegantice-
ras (Lehmann 1967). In addition, incompletely
preserved remnants of radulae have been recor-
ded in a few genera. In our material of the Ap-
tian ammonite Aconeceras, about twenty juvenile
shells have radula, aptychus and counter-apty-
chus preserved in the living chamber, The radula
is composed of numerous teeth-rows (r, pl. 1,
Figs. 2, 3). Each row has seven teeth (Fig. 2c) :
the rachidial tooth (R) with five to seven cusps ;
the two lateral teeth (L1, L2) with five cusps
each; the extremely high, curved and cone-shaped
marginal tooth (M1 ; see also pl. 1, Fig. 3) ; and
the small marginal plate (MP). The radula teeth
seem to be at least partially calcified. The radula
of Aconeceras has the same number of teeth in

PLATE 1
Aconeceras.
Fig. 1 - Juvenile shell in lateral view, about 15 mm in diameter and consisting of four whorls, to show the
aptychus (a) in the living chamber (lch). Vue lat$rale d'une coquille juvdnile (environ 15 mrn de diarn~tre et
quatre tours), montrant l'aptychus (a) dans la loge d'habitation (lch).
Fig. 2 - Longitudinal section of the same aptychus (a) to show rows of radular teeth (r), x 50. Section longitudinale
du rhyme aptychus (a) montrant les rangdes de dents radulaires (r).
Fig. 3 - Detail of the same radula ; note the long and curved marginal teeth (M), x 150. Ddtail de la rn~me radula ;
noter la dent latdrale longue et courbe.
Geobios PI. 1
M.S. n" 15 L.A. D o g u z h a e v a & H. M u t v e i
 116
each row as those in E o a s i a n i t e s a n d E l e g a n t i c e -
ras, but differs in the shape of the teeth : in
E o a s i a n i t e s the teeth are all uniformly long and
cone-shaped, whereas in E l e g a n t i c e r a s t h e t w o la-
teral teeth are cone-shaped and the three middle
teeth each have three cusps. The r a d u l a in
A c o n e c e r a s can be classified with the ctenoclossal
type, being extremely similar to the radula in in-
cirrate octopods (compare Fig. 2c,d). This m a y in-
dicate a phylogenetic relationship between the in-
cirrate octopods and ammonoids.
The radulae (r) are surrounded by the valves of
aptychi and in certain cases even by counter-ap-
tychi (a, ca, Fig. 2 a ; a, pl. 1, figs. 1, 2, 3). Our
preliminary studies show t h a t the anterior edge
of the aptychial valves is completely calcified and
composed of numerous consecutive prismatic sub-
layers (ac, Fig. 2 a,b). Apart of the anterior calci-
fied edge, the aptychial valves axe composed of
two layers : an outer mainly organic layer and an
inner mainly calcified layer (ol, cl, Fig. 2b). The
outer organic layer is composed of numerous thin
lamellae and it seems to be only slightly calcified.
e i:;i
f ,i"
c~
Figure 3 - Ptychoceras. a : median section of a species with
untruncated initial portion of the shell, b : detail of the initial
portion of the shell ; note that the dorsal shell wall (dwl) is
covered by the dorsal wall of the second, shaft; (dw2). r : me-
dian section of a species with truncated initial portion of the
shell, d, e : detail of the truncated end of the first shatt in
two shells ; note that the truncated end became enclosed by
the dorsal walls of the second and third shaits (dw2, dw3) at
later ontogenetic stages, a : section longitudinale d'une estate
avec partie initiale, non tronqude, de la coquille ; b : dgtail de
la pattie initiale de la coquille ; lernur dorsal (dwl) est couvert
par le tour dorsal de la seconde gaine ; e : section longitudi-
nale d'une esp~ce dont la pattie initiale e s t tronquge ; d, e :
details de l'extrgmit~ tronqu~e de la premiere gaine sur deux
coquilles ; il est a noter que la pattie tronquffe est entourde par
les murs dorsaux des seconde et troisi~rne gaine (dw2, dw3).
The inner calcified layer in places shows a pris-
matic structure, but this structure is probably
usually obliterated by a high content of organic
matrix, The inner lamella is short and thin (il,
Fig. 2b).
The counter-aptychi also seem to be composed of
two valves which have a structure similar to t h a t
in aptychi : the anterior edge is completely calci-
fied and the valves are slightly calcified (ca, ac,
Fig. 2a). However, the valves of the counter-apty-
chi, including the anterior calcified edge, are
much t h i n n e r t h a n those in aptychi, and there-
fore less well preserved. Details of the structure
in counter-aptychi are still not fully understood
and will be dealt with in a forthcoming paper.
SHELL TRUNCATION IN
HETEROMORPHIC PTYCHOCERAS
Shell structure was studied in four species of the
heteromorphic lytoceratid genus P t y c h o c e r a s from
the Aptian of N. Caucasus (Doguzhaeva &
 117

F i g u r e 4 - P t y c h o e e r a s . H y p o t h e t i c a l r e c o n s t r u c t i o n of t h e
r e l a t i o n s h i p b e t w e e n t h e soft body a n d s h e l l : in a : t h e s h e l l
is r e c o n s t r u c t e d a s i n t e r n a l , a n d i n b : a s s e m i - i n t e r u a l . Re-
construction hypoth$tique des relations entre la pattie molle
des corps et la coquille ; a : la coquille est supposde interne ; b:
elle est supposde semi-interne.

Mutvei 1989). The shell is composed of an initial with g). The Triassic ceratid Megaphyllites is par-
planispiral whorl and three parallel, straight ticularly suitable for studies on the mode of tran-
shales (Figs. 3 a,b). In three of these four species sition from retrochoanitic to prochoanitic septal
the shell was truncated during the lifetime, i.e. neck, because the transition took place at an on-
its initial coiled portion, comprising about fifteen togenetically late stage, and because the site of
chambers, was broken off (Fig. 3c). After the transition is long, comprising about two whorls
truncation, the broken end of the shell was lei~ (Fig. 5a ; Doguzhaeva & Mutvei 1986b). However,
open until it became embedded between the se- it must be emphasized that this transition occur-
cond and the third shales (dw2, dw3, Fig. 3 d,e). red only in the dorsal side of the siphonal tube,
whereas the ventral side remained retrochoanitic
The shell wall of Ptychoceras has a unique and (Fig. 5 a,b,c,d). The transition began in such a
complex structure. The outer shell surface, at way that the septum increased in thickness at
least in the apertural region, is covered by a thin the passage to the septal neck and formed a fold
external nacreous coating-layer which seems to (sf, Fig. 5c) which increased in height in sub-
have been secreted from the outside. Many shells sequent septa. At the final stage of development,
show fragmentation and repair of the shell wall. the septal fold was subdivided into two halves :
The shell's struncation, its unique structure, the the inner half with the retrochoanitic neck for-
secretion of the nacreous coating-layer, and the med the cuff, whereas the outer half became the
fragmentation and repair of the shell, indicate prochoanitic septal neck (cuff, pn, Fig. 5b).
that the shell has been either partially (Fig. 4b)
or entirely (Fig. 4a) covered by the mantle. In most adult ammonites, the siphonal tube is
fully prochoanitic (pn) both on the ventral and
dorsal sides, The cuffs are not nacreous, as in
TRANSITION RETROCHOANITIC/ Megaphyllites, but show a prismatic structure
PROCHOANITIC SEPTAL NECKS (cuff, Fig. 5g).
A common evolutionary trend in ammonoids is During evolution, the appearance of the prochoa-
the replacement of the retrochoanitic septal nitic septal necks shii~ed successively to earlier
necks by the prochoanitic necks (compare Fig. 5f ontogenetic stages. So, in most ammonitids, the
 118

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Figure 5 - a-d : transition of retrochoanitic necks (rn) to prochoanitic necks (pn) in the second (II), third (III) and fourth whorl
(IV) in the ceratid M e g a p h y l l i t e s ; a : general view ; b, c, d : siphonal tube in the fourth, third and second whorl, respectively ;
note that the retrochoanitic neck (rn) is retained as the cuff. e : retrochoanitic s~phonal tube in N a u t i l u s , and f : in the goniatid
A g a t h i c e r a s ; note that the connecting ring in the former has two layers (sph, ol) but in the latter one layer (ol) ; g : prochoani-
tic siphonal tube (pn) in the lytoceratid G a u d r y c e r a s to show the modified cuff = remnant of the retrochoanitic neck. Structures
siphonales, a-d : transition depuis les goulots rdtrochoands (rn) aux goulots prochoands (pn) dans le second (II), le troisi~me (III)
et le quatri~me (IV) tour chez le Ceratitidd M e g a p h y l l i t e s ; a : vue gJndrale ; b, c, d : tube siphonal, respectivement sur les tours
IV, III, H ; le goulot retrochoan~ est conserv~ comme manchon, e, f : tube slphonal retrochoan~ du N a u t i l u s (e) et de la goniatite
A g a t h l c e r a s ; l'anneau connectif poss~de deux couches (sph, ol) sur le premier reals une seule sur le second (ol) ; g : tube siphon~l
pochoanJ (pn) du lytoceratid~ G a u d r T c e r a s montrant le manchon rnodifid qui est un reste du goulot retrochoan&

r e t r o c h o a n i t i c s e p t a l n e c k s do n o t occur, b u t t h e
p r o c h o a n i t i c n e c k s w e r e f o r m e d f r o m t h e begin-
n i n g of ontogeny. M o r e o v e r , d u r i n g evolution, t h e
site of t r a n s i t i o n f r o m r e t r o c h o a n i t i c to p r o c h o a -
nitic s e p t a l n e c k s b e c a m e s u c c e s s i v e l y s h o r t e r .
T h e l e n g t h of t h e t r a n s i t i o n site is d i f f e r e n t in
different phylogenetic lineages (Doguzhaeva &
M u t v e i 1986b).
CONCLUDING REMARKS
T h e r e s u l t s of o u r s t u d i e s c o n f i r m t h a t p r i n c i p a l
a n a t o m i c a l differences e x i s t e d b e t w e e n a m m o -
noids a n d t h e R e c e n t N a u t i l u s in m u s c l e s y s t e m ,
buccal a p p a r a t u s , s h a p e o f h e a d - f u n n e l r e g i o n
a n d its r e l a t i o n s h i p to shell a p e r t u r e , e x t e n s i o n
of m a n t l e to shell surface, a n d f u n c t i o n a n d
s t r u c t u r e o f s i p h o n a l tube. On t h e o t h e r h a n d ,
certain similarities were found between amino-
h o l d s a n d R e c e n t coteoids : (1) a s i g n i f i c a n t simi-
laxity in s h a p e of r a d u l a r t e e t h w i t h b a t h y p e l a g i c
octopods, (2) t e n d e n c y to d e v e l o p a n i n t e r n a l
shell, a n d (3) possible d e v e l o p m e n t o f s t r o n g hy-
ponome retractors.
A c k n o w l e d g e m e n t - This study was financially sup-
ported by Grants 287-113-116 of the Swedish Natural
Science Research Council.
 119
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