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Abstract
Palaeontological Institute of the Russian Lebedev, O.A. 2009. A new specimen of Helicoprion Karpinsky, 1899 from
Academy of Sciences, Profsoyuznaya St. Kazakhstanian Cisurals and a new reconstruction of its tooth whorl position
123, Moscow, 117997, Russia and function. — Acta Zoologica (Stockholm) 90 (Suppl. 1): 171–182
A new Helicoprion bessonowi Karpinsky, 1899 (Chondrichthyes, Eugeneodonti-
Keywords:
Lower Permian, Artinskian, Helicoprion, formes) specimen from the Artinskian of Kazakhstan is described. This is the
dentition southernmost occurrence of this species in the Cisurals area. Its presence
suggests a biogeographical link for this species between the Cisurals and Japan.
Accepted for publication: Residue obtained from chemical preparation of the sample included numerous
18 June 2008 scales and several teeth, which are tentatively assigned to Helicoprion. This
assumption is based upon morphological similarity of the scales to those
known in other eugeneodontiforms. Campodus-like teeth might be part of the
lateral dentition of Helicoprion. A new reconstruction of the interaction of the
lower tooth whorl with the upper jaw dentition is suggested and its function is
discussed. It is proposed that there was no symphysial whorl in the upper jaw
but its role was played by a rigid cover formed by a series of small teeth at the
palatoquadrates. Microscopic study of the tooth crown surface revealed scratch
marks, which might have resulted from pressing the food object against the
upper jaw. Using extant odontocetans as an ecological model led to a conclusion
that helicoprionids most likely fed on cephalopods and to some extent on fish.
This assumption is based upon the concentration of functional dentition in the
area of the lower jaw symphysis in both groups of animals.
Oleg A. Lebedev, Palaeontological Institute of the Russian Academy of Sciences,
Profsoyuznaya St. 123, Moscow, 117997, Russia.
E-mail: olebed@paleo.ru
et al. 2000), Japan (Yabe 1903), Laos (Hoffet 1933), China Stethacanthus sp. (Fig. 4N–P) and scales of Actinopterygii indet.
(Chen et al. 2007), Australia (Teichert 1940) and possibly (Fig. 4Q ). These will be described elsewhere. Microremains
Iran (Obruchev 1964). The youngest Helicoprion find comes were examined by scanning electron microscopy.
from the Lower Kungurian of the Perm Region (Tchuvashov
2001). Several finds of Sinohelicoprion (Liu and Chang 1963;
Systematic palaeontology
Zhang 1979; Liu and Wang 1994), including the latest one
from the Changxingian of Hunan province (Lei 1983) and Order Eugeneodontiformes Zangerl, 1981
Hunanohelicoprion (Liu 1994; Cheng et al. 2004) including Family Helicoprionidae Karpinsky, 1911
the youngest species from the Maokuo of the Guizhou prov-
ince (Cheng et al. 2004), are known from China. Therefore,
Referred genera
helicoprionids were globally distributed apart from the
southwest margins of Gondwana. This may be the result of Apart from the type genus, Agassizodus St John and Worthen,
supposed glaciation preventing their spreading to this region 1875; Parahelicoprion Karpinsky, 1924; Campyloprion East-
during the Early Permian. These huge sharks possibly inhabited man, 1902; Sarcoprion Nielsen, 1952; Toxoprion Hay, 1909;
the oceanic basins and entered the shallow-water seas only Sinohelicoprion Liu and Chang, 1963; Hunanohelicoprion Liu,
temporarily, for example during spawning seasons. 1994 and Shaktauites Tchuvashov, 2001.
The present finding represents the southernmost
occurrence of Helicoprion bessonowi Karpinsky, 1899 in the
Diagnosis (emended after Obruchev 1964)
early Permian Cisurals basin (Fig. 1). The new locality of
H. bessonowi Karpinsky, 1899 is situated at the junction of Large marine eugeneodontiforms in which the jaw symphysis
the Uralian Strait with the northern coast of Tethys, providing was armed with an arched or spiral dental complex composed
a biogeographic connection to the occurrence of this species of the conjoint base and a set of separate tooth crowns
in Japan as well as evidence of a Tethys–Panthalassa link compressed laterally. Lateral crown sides form symmetric
between Asian and Euramerican helicoprionids. processes extended rostrally (spurs), whereas the bases
(where seen) extend caudally. Lateral teeth organized into
numerous series, up to 25 elements in each. The base of the
Materials and Methods
tooth whorl rests upon an unpaired spiral basimandibular
A new H. bessonowi Karpinsky, 1899 (specimen PIN 1988/38, rod-like cartilage fusing with the Meckelian cartilages.
Fig. 2) was found by Ruzhentsev in Kazakhstan (Aktyubinsk
Region, Aktasty locality; Lower Permian, Artinskian, Aktasty
Remarks
Substage). The specimen consists of two almost half-volutions,
the larger one from the youngest, external coil and the smaller The family Helicoprionidae was created by Karpinsky as
one from the inner coil adjoining it. Both were embedded in early as 1911, but apart from the Russian palaeoichthyologists
a piece of organoclastic limestone. The external face of the this systematic unit was accepted only by Teichert (1940)
specimen was badly damaged.To expose the unbroken internal and Bendix-Almgreen (1966). Most western researchers
surface of the whorl the sample was treated with 10% acetic (for example Chorn 1978) included this genus into the
acid solution. The residue obtained comprised numerous Edestidae until Zangerl (1981) erected the Agassizodontidae
diverse microremains, including scales and teeth.The associated to include genera related to Helicoprion Karpinsky, 1899. He
fish microremains included a variety of other chondrichthyan noted partial correspondence of the Agassizodontidae to
teeth like those of Cobelodus obliquus Ivanov, 2005 (Fig. 4K–M), Helicoprionidae but did not explain the reasons for erecting
Fig. 2—Two fragments of the tooth whorl PIN 1988/38, Kazakhstan, Aktyubinsk Region, left bank of the Aktasty River, Aktasty locality;
Lower Permian, Artinskian, Aktasty Substage. Scale bar = 3 cm.
a new family name. As the Helicoprionidae Karpinsky are known: Helicoprion sierrensis Wheeler, 1939 (Asselian-
(1911) have priority over Agassizodontidae the former family Artinskian, glacial boulder, California); H. nevadensis Wheeler,
name is supported here. 1939 (Artinskian, Nevada); H. ferrieri (Hay, 1907) and H.
ergasaminon Bendix-Almgreen, 1966 (Phosphoria Formation,
Helicoprion Karpinsky (1899a) Roadian; Idaho); H. svalis Siedlecki, 1970 (Artinskian-
Diagnosis (emended after Obruchev 1964 and Zangerl 1981) Ufimian; Spitsbergen); H. davisii (Woodward, 1886) (?
Kungurian-Ufimian; Western Australia); H. mexicanus
High and narrow symphysial part of the Meckelian cartilage Müllerried, 1945 (Leonardian, Mexico). Poorly preserved
forms a case enclosing most of the older part of the spiral. H. karpinskii Obruchev, 1953 specimen comes from pre-
Symphysial dentition forms a whorl, which may consist of up sumably Kungurian of Cisurals.
to three coils and bear up to 180 tooth crowns. Crown cutting
edges smooth or denticulated. Lateral processes of crowns Helicoprion bessonowi Karpinsky (1899a) (Figs 2 and 3)
(spurs) attain the level of the vertical axis of the preceding or
the next before the preceding tooth crown rostrally. Teeth Helicoprion bessonowi: Karpinsky (1899a; p. 20, text-figs 18 –
consist of osteodentine and the crowns are coated with 57, pls. 1–3, 4, figs 1–11); Karpinsky (1899b; p. 25, text-
enameloid. Palatoquadrates are lined with the series of small figs 18–22 and 36–57); Karpinsky (1911; p. 1105, text-
‘Campodus’-like teeth. fig. 2); Obruchev (1964; pl. 3, fig. 1, text-figs 32–33);
Zangerl (1981; p. 86, text-fig. 98I).
Type species
Helicoprion bessonowi Karpinsky (1899a). Type specimen
Geographical distribution and age Lectotype – tooth whorl in the Central Research Geological
Museum (TsNIGR, Russia, St.-Petersburg) 1/1865, Russia,
Apart from the type species from the Artinskian of Cisurals Sverdlovsk Region, Krasnoufimsk District, a quarry 3 km to
(Russia), Kazakhstan and Japan, the following species the west from Krasnoufimsk; Lower Permian, Artinskian,
transversely and extends to the lingual side of the crown more rectangular or rhombic teeth disposed in rows in the Helico-
than to the labial one. Two pairs of lateral tubercles bear a prion ferrieri specimen of the anterior part of the head. Both
low, but acute sagittal ridge, often giving off small branches. upper and lower jaws in Sarcoprion (Nielsen 1932, 1952)
At the central tubercle the sagittal ridge is interrupted by bear numerous rows of Campodus-like teeth. It is highly
a transverse crest. Short crests fill the space between the probable that the tooth PIN 1988/47 was a part of Helicoprion
branching ornament of the main system of ridges.This specimen lateral dentition.
is morphologically indistinguishable from the one figured by
Karpinsky (1904), but is much smaller. ? Helicoprion bessonowi Karpinsky (1899a) (Figs 4C–J and
5M–P)
Remarks
Description
Remains associated with the Helicoprion tooth whorls are
very rare. Karpinsky (1904) presented an unassociated Numerous scales extracted from the rock sample are uniformly
Campodus sp. tooth originating from the same locality from built, although they show a great variety of individual appearance.
where an imprint of the whorl was known. He suggested, and Generally a high coronal part is mounted upon a shallow
later Nielsen (1952) supported the suggestion by analogy rounded, oval or irregularly rounded base forming a narrow
with Sarcoprion, that this tooth belonged to Helicoprion lateral brim around the crown footing. The basal surface is concave,
dentition. Bendix-Almgreen (1966) described minute irregular and bears numerous vascular pores (Fig. 4H).
Fig. 5—Comparison of morphological scale patterns in various eugeneodontiforms. —A. Scales at the lateral side of the lower jaw symphysis
in Sarcoprion edax Nielsen, 1952 (large elements on top of the figure are basal parts of the tooth whorl crowns); B–H. ‘edestid’ scales from
the Upper Permian of East Greenland; —I–L. scales in Ornithoprion hertwigi Zangerl, 1966, all in vertical cross-section; M–P. scales
presumably belonging to Helicoprion bessonowi Karpinsky, 1899. —M. Specimen PIN 1988/51; —N. specimen PIN 1988/48; —O. specimen
PIN 1988/50; —P. specimen PIN 1988/49. —A, C, E, P. coronally; —B, D, F, G, H, M–O. laterally. —A. redrawn from Nielsen (1952);
—B–H. redrawn from Stensïo (1962); —I–L. from Zangerl (1966), modified. Not to scale.
Monodontode or polyodontode crowns are sculptured by (Fig. 5). All of them demonstrate a flat or somewhat concave
rough vertical ridges, which tend to become finer and bifurcate base rounded in the plan view and generally following the
basally. The ridge margins are serrated (Fig. 4I). In poly- crown outline. The neck is better expressed in the elements
odontode scales individual conical odontodes fuse with their consisting of a smaller number of odontodes (Fig. 5F,G,I,K,O).
bases and basal parts of their walls, but their apical parts The crown is formed by a differing amount of odontodes
remain independent (Fig. 4C–F). In monodontode scales a fused at their sides. Individual odontode apices are expressed
slender cusp occupies a small area in the middle of the wide to varying degrees, being rounded in Ornithoprion (Fig. 5I – L)
flat base (Fig. 4J). and Greenland ‘edestids’ (Fig. 5B – H), pointed in the
Helicoprion bessonowi scales (Fig. 5M–O) and flat (not
unlikely because of lifetime abrasion) in Sarcoprion
Remarks
(Fig. 5A). This general morphological similarity between
Scales were never found in association with Helicoprion scales in the edestiforms mentioned above does not make
whorls, even with the best-preserved specimens of H. ferrieri improbable the attribution of the materials from Kazakhstan
(Hay 1907). Bendix-Almgreen (1966), who described this to Helicoprion.
material in detail, noted that this was probably because of the An alternative suggestion is the attribution of these elements
absence of scales at the snout. However, in Sarcoprion, to other fish, for example symmoriiforms Cobelodus and
another member of the Helicoprionidae, Nielsen (1932, Stethacanthus. However, their skin is naked except for modified
1952) figured the lateral part of the symphysial region lined scales along the lateral line canal (e.g. Zangerl 1981), so
with scales possessing flattened polygonal crowns (Fig. 5A). scales described above might belong to either an unknown
Presumably ‘edestid’ scales from the Upper Permian of East fish of which there are no other remains in the collection,
Greenland (possibly belonging to Fadenia or Sarcoprion) or to Helicoprion, which is easier to assume. The second
were described by Stensïo (1962) (Fig. 5B– H). In eugeneo- interpretation seems more parsimonious and is accepted
dontiform caseodontid Ornithoprion from the Pennsylvanian here.
of Indiana, USA (Zangerl 1966) scales are known (Fig. 5I–L).
Taking into account manifestation of these skeletal elements
Discussion
in related members of the order, one cannot exclude the
possibility of their presence in Helicoprion. Various authors suggested several hypotheses on the position
Morphologically, scales in Ornithoprion, East Greenland of the Helicoprion tooth whorl and its function. Russian workers
‘edestids’ and those described above are very closely built (Karpinsky 1899a,b; Obruchev 1953, 1964; Tchuvashov
2001) placed the whorl at the upper jaw and this reconstruc- anterior part of the external coil exposed out of the mouth
tion remained in textbooks for decades (Fig. 6). Karpinsky cavity, representing the rostral and ethmoidal parts of the
(1911, 1915) suggested that the whorl might be enclosed in skull as being very low and hypothesized that the whorl
the cartilage, but his first earlier graphic reconstruction did rested between ‘two similar series of upper teeth’. On the
not show this (Fig. 6A). Obruchev (1953) reconstructed the contrary, Bendix-Almgreen (1966) concluded that ‘... no
whorl with the symphysial cartilage filling the space between similar organ was developed in the upper jaw’.
the coils. His idea was based upon the presence of prismatic Apart from Sarcoprion, whorls in both the lower and the
cartilage patches remaining there after preparation (Fig. 6B). upper jaws within Eugeneodontiformes are known only in
It is not improbable that some of the whorls were preserved Fadenia crenulata Nielsen, 1932 (Bendix-Almgreen 1975),
in association with cartilage enclosing them but it was later Edestus heinrichi Newberry and Worthen, 1870 (Zangerl
removed by careless preparation. 1981) and Edestus mirus Hay, 1912 (Nielsen 1952). In the
Bendix-Almgreen (1966), on the basis of uniquely pre- last two forms, the position of these elements with respect to
served materials from the Phosphoria Formation (Idaho) each other is either unknown or dubious. Superposition of
clearly stated that the Helicoprion whorls were set at the lower the Fadenia lower jaw to the upper one suggests that the
jaw symphysis, rested upon an unpaired basimandibular former was somewhat shifted caudally, so that occlusion of
cartilage and were enclosed, apart from the youngest part of the anterior lower jaw elements possibly occurred with the
the external coil, into a cartilaginous case formed by the posterior upper jaw elements. Moreover, in Sarcoprion and
symphysial part of the Meckelian cartilages. Regretfully, he Fadenia symphysial elements are much wider and, respec-
reconstructed only this anteriormost part of the lower jaw tively, lower, more robust than in Helicoprion, which makes
and did not demonstrate its position with respect to the upper food processing between whorls conceivable, in contrast to
jaws and rostrum. The first author to publish the reconstruction the last genus in which dental crowns are very high, strongly
of the tooth whorl at the lower jaw in the whole fish was Long compressed laterally and would hardly resist lateral force
(1995) (Fig. 7A). Janvier (1996) also placed the whorl at the which would inevitably arise from interaction with the upper
lower jaw and enclosed half of it within a reconstructed jaw symphysial antagonist. Taking into account dozens,
cartilaginous case (Fig. 7B). However, the latter author was maybe hundreds, of Helicoprion tooth whorls found in the
following the structure of the lower and upper jaws of the world, not a single ‘upper jaw’ specimen was ever described.
related long-snouted genus Sarcoprion. Janvier displayed the No upper jaw element was found in the best-preserved specimen
less complete skeletons (Caseodus, Romerodus and Fadenia: makes about 25% of the skull length, in Caseodus about 28%
Zangerl, 1981) the body is fusiform, the caudal fin is equilobate and in Fadenia about 40%. This range of proportions is likely
and strongly forked, suggesting outstanding swimming to be close to the meaning in Helicoprion. The largest known
abilities, as should be also the case in Helicoprion. Only one whorl diameter is 0.4 m (Tchuvashov 2001), which would
dorsal fin appears to be present. Neither anal nor pelvic fins correspond to a 1.0–1.6 m skull length and, correspondingly
are reconstructed by analogy with its closest eugeneodon- to a 5–8 m body length, which is comparable to the length of
tiform relatives. the largest modern sharks. Certainly, this estimate is very rough
Spectators encountering the Helicoprion tooth whorls are and is based upon assumptions, which cannot currently be
always impressed by their large size. However, no attempts checked.
to guess the total length of the body were ever made by
palaeoichthyologists. It is suggested here to use data on the
Conclusions
whorl size for at least very rough calculation of this parameter.
If the whorl diameter is roughly equal to the length of the lower – A new find of Helicoprion bessonowi Karpinsky, 1899
jaw symphysis, it may be used for an approximate estimation enlarges the distribution of these fish to the South Urals
of the skull length. Using this standard, and suggesting ‘normal’ area, which corresponded during the Permian to the
fish body proportions (that is not too much elongated eel- northern coast of Tethys, providing a biogeographical
like or shortened) in Helicoprion, the body length should be connection to the occurrence of this species in Japan
around five times the head length, that would give us very through the Uralian Strait.
rough estimate of the body length. Thus, basing upon illu- – Simple monodontode and complex polyodontode scales
strations in Zangerl (1981), in Romerodus the symphysis length found in the rock sample probably belong to Helicoprion
based upon morphological similarity to those known in Hay, O. P. 1907. A new genus and species of fossil shark related to
other eugeneodontiforms. Campodus-like teeth might be Edestus Leidy. – Science 26: 22–24.
a part of Helicoprion lateral dentition. Hay, O. P. 1909. On the nature of Edestus and related genera, with
descriptions of one new genus and three new species. – Proceedings
– A new reconstruction of the symphysial tooth whorl
of the United States National Museum 37: 43–61.
contact with the snout and upper jaw structures and Hay, O. P. 1912. American Permian vertebrates. – American
whorl function is proposed and the whorl function is Naturalist 46: 561–565.
discussed. The younger third of the last coil is thought to Hoffet, J. H. 1933. Ètude géologique sur le centre de l’Indochine
enter a median palatal cavity. It is proposed that there entre Tourane et le Mekong. – Bulletin de la Service géologique du
were no analogous whorl antagonists in the upper jaw but Indochine 20: 3–154.
Ivanov, A. 2005. Early Permian chondrichthyans of the Middle and
its role was played by rigid cover formed by series of small
South Urals. – Revista Brasileira de Paleontologia 8: 127–138.
teeth on the palatoquadrates. Janvier, P.H. 1996. EarlyVertebrates, pp. 5 – 393. Clarendon Press, Oxford.
– Microscopic study of the tooth surfaces revealed scratch Karpinsky, A. P. 1899a. On the edestid remains and its new genus
marks which might have resulted from pressing of the food Helicoprion. – Zapiski Imperatorskoy Akademii Nauk 7: 1– 67 (in
object against the upper jaw, rather than from ploughing Russian).
bottom sediment in search for soft invertebrates as was Karpinsky, A. P. 1899b. Über die Reste von Edestiden und die neue
suggested by earlier authors. Gattung Helicoprion. – Verhandlungen Russisches Kaiserliches
Mineralogisches Gesellschaft 36: 361– 475.
– Using extant odontocetans as a morphonutritiological
Karpinsky, A. P. 1904. Presence of the remains of the genus
model leads to a conclusion that helicoprionids most Campodus de Koninck in the Artinskian deposits of Russia. –
likely fed on cephalopods and to some extent on fish. This Zapiski Imperatorskogo Sankt-Peterburgskogo Mineralogicheskogo
assumption is based upon concentration of dentition at Obshchestva 41: 32–37 (in Russian).
the lower jaw symphysis in both groups of animals. Karpinsky, A. P. 1911. Notes on Helicoprion and other edestids. –
Izvestiya Imperatorskoy Akademii Nauk 5: 1105–1122 (in Russian).
Karpinsky, A. P. 1915. On the nature of the spiral organ of Helicoprion.
Acknowledgements – Zapiski Ural’skogo Obshchestva Lyubiteley Estestvoznaniya 35:
117–145 (in Russian).
The author would like to express his sincere gratitude to Karpinsky, A. P. 1924. Helicoprion (Parahelicoprion n.g.) clerci. – Zapiski
Prof. Janvier (MNHN, Paris, France) and other colleagues – Ural’skogo Obshchestva Estestvoispytatelei 34: 1–10 (in Russian).
palaeoichthyologists – for encouraging this study and for pro- Kelly, M. A. and Zangerl, R. 1976. Helicoprion (Edestidae) in the
viding valuable comments during his report in Uppsala Permian of west Texas. – Journal of Paleontology 50: 992–994.
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Late Permian of Hunan, South China. – Vertebrata Palasiatica 21:
manuscript preparation were supplied by anonymous referees,
347–351.
to whom the author is greatly indebted. Liu, Z.-H. 1994. New material of helicoprionid shark from Lianyuan
of Hunan. – Vertebrata Palasiatica 32: 127–133.
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