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«Ann. Naturhist. Mus. Wien Wien, Februar 1990 53-61 A Neogene Cobras of the Genus Naja (Serpentes: Elapidae) of East Europe Neogene Kobras der Gattung ...»

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©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at

Ann. Naturhist. Mus. Wien Wien, Februar 1990



Neogene Cobras of the Genus Naja (Serpentes: Elapidae)

of East Europe

Neogene Kobras der Gattung Naja (Serpentes: Elapidae) aus Osteuropa


(Mit 3 Abbildungen im Text)

Manuskript eingelangt am 18. September 1989


Fossile Überreste der Gattung Naja sind aus drei neogenen Fundstellen bekannt: Gritsev (Ukraine), Kohfidisch (Österreich) und Tourkobounia 1 (Griechenland). Reste aus den beiden ersterwähnten Fundstellen gehören der ausgestorbenen Art Naja romani an, die ursprünglich aus dem Miozän Frankreichs beschrieben wurde. Diese Art bildet einen frühen Ausläufer der Entwicklungslinie, die zu den rezenten asiatischen Vertretern der Gattung Naja führt. Aus dem Vorkommen von N. romani in den letzten sechs Millionen Jahren gehen neue Erkenntnisse über die Langlebigkeit der miozänen Ophiden-Arten hervor.

Summary Fossil remains of the genus Naja are reported from three Neogene localities: Gritsev (Ukraine), Kohfidisch (Austria), and Tourkobounia 1 (Greece). Remains from the two former localities belonged to the extinct species Naja romani originally described from the Miocene of France. This species represented an early offshoot of the evolutionary lineage leading to the living Asiatic members of the genus Naja. The persistence of N. romani for at least six million years provides new evidence for the longevity of Miocene ophidian species.

Introduction Among extinct European snakes, the Elapidae have perhaps the most well documented history. The fossil record of the family, which at present does not inhabit the discussed area, comes from fifteen Neogene localities. Most of the excavated material consists of isolated vertebrae only, but the cranial elements -, usually well preserved, also occur quite commonly. SZYNDLAR and RAGE (in press)

Anschrift der Verfasser:

') Dr. Zbigniew SZYNDLAR, Institute of Systematic and Experimental Zoology, Polish Academy of Sciences, Slawkowska 17, PL-31-016 Krakow. - Poland.

) Mrs. Galina A. ZEROVA, Department of Paleozoology, Institute of Zoology, Ukrainian Academy of Sciences, Lenina 15, SU-252650 Kiev-30. - USSR.

©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at 54 Z. SZYNDLAR & G. A. ZERO VA recently reviewed the entire elapid fossil record from Europe and adjacent territories and outlined phylogenetic interrelationships among both extinct and extant cobras. They demonstrated that all but one of the European elapids belonged to the modern genus Naja. The latter research confirmed SZYNDLAR'S (1985) opinion that living members of the genus Naja represented two separate phyletic lineages, currently inhabiting Africa and Asia, respectively. In the past, members of the former complex occurred in the Iberian Peninsula, while the remaining part of southern Europe was occupied by representatives of the Asiatic lineage. Most finds of European cobras come from western Europe; in the present paper we review Neogene fossils found in three East European sites in Ukraine, Austria, and Greece.

A c k n o w l e d g e m e n t s. - We are grateful to the late Professor Friedrich BACHMAYER and Dr.

Ortwin SCHULTZ (Naturhistorisches Museum, Vienna), and to Dr. Hans de BRUIJN (Rijksuniversiteit Utrecht) for loan of the fossil material to study. We thank Dr. Aaron M. BAUER (Villanova University) for his valuable criticism and advice. Visits of the senior author in Vienna (1988), Kiev (1989), and Paris (1989) were possible thanks to grants of the Naturhistorisches Museum in Vienna, Centre National de la Recherche Scientifique of France, and Academy of Sciences of the USSR.

We wish to dedicate this short article to the memory of Professor BACHMAYER in order to honour his prominent contribution to our knowledge on the Kohfidisch fauna.

Localities and material The oldest locality of Gritsev is located in Khmelnitsky oblast', western Ukrainian SSR. According to KOROTKEVICH (1988), its fauna is of late Miocene age (Neogene European Land Mammalian Zone MN 9, Mammalian Age lower Vallesian). The squamate fauna from Gritsev was preliminary listed by ZERO VA (1987), and an erycine snake {Albaneryx) was described in detail by the same author (ZEROVA 1989). Snake remains belong to the following taxa: an unidentified scolecophidian, Albaneryx volynicus and Bransateryx sp. (Boidae: Erycinae), a natricine, Coluber sp., Elaphe sp. (Colubridae), Vipera sp. (Viperidae), and a member of the genus Naja (Elapidae). The latter snake was identified on the basis of a basisphenoid, a right compound bone, and ten trunk vertebrae. The material is the property of the Institute of Zoology of the Academy of Sciences of Ukrainian SSR (IZAN).

Kohfidisch, located in Burgenland (Austria), is of late Miocene age (MN 11, lower Turolian, after BACHMAYER & ZAPFE 1969). The snake fauna, described in

detail by BACHMAYER & SZYNDLAR (1985, 1987), consists of five extinct species:

Elaphe kohfidischi, Coluber planicarinatus, Natrix longivertebrata (Colubridae), Vipera burgenlandica (Viperidae), and Naja austriaca (Elapidae).The description of Naja austriaca was based on a basisphenoid (holotype) and a set of other cranial and axial bones. The newly discovered material, described in the present paper, contains two other basisphenoids, a fragmentary parietal, one right and one left prootics, one basioccipital, two squamosals, two left quadrates, ribs, and vertebrae. The material belongs to the Naturhistorisches Museum in Vienna (NMW).

©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at

–  –  –

Fig. 1: Basisphenoids of Naja romani from Kohfidisch (A-C and D-F; NMW 1989/34/1 and 1989/34/2) and from Gritsev (G-I; IZAN 22-1771); ventral (A, D, G), left lateral (B, E, H), and dorsal (C, F, I) views. Scale line 5 mm. Abbreviations: aovc, anterior orifice of Vidian canal; povc, posterior orifice of Vidian canal; V4, groove and foramina for entry and exit of cid-nerve; VI, foramina for abducens nerve.

©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at 56 Z. SZYNDLAR & G. A. ZEROVA Tourkobounia 1, lying in the area of Athens (Greece), is of upper Pliocene age (MN 16, Villanyian; de BRUIJN & van der MEULEN 1979). The snake fauna is comprised of a scolecophidian, Malpolon sp. (Colubridae), two species of Vipera (Viperidae), and Naja sp. (Elapidae), the latter represented by seventeen trunk vertebrae. The material belongs to the Geological Institute of the Utrecht University (UUGI).

Morphological description In the following description we focus on elements having taxonomic importance; the bones for which morphology is similar in all or most species of Naja are not described in detail.

The three available basisphenoids, both those from Kohfidisch and that from Gritsev (Text-fig. 1), display the same morphological patterns and do not differ from the holotype basisphenoid of Naja austriaca (cf. BACHMAYER & SZYNDLAR 1985: 89, Text-fig. 3). The parasphenoid portion is missing in all the bones. The basisphenoid portion is hexagonal in shape and becomes distinctly narrower immediately before the basipterygoid processes; the latter structures are strongly developed. The posterior orifices of the Vidian canals are located on the lateral margins of the bones (i. e., at the suture between the basisphenoid and prootic in a complete braincase). The anterior orifices of the canals lay on the ventral side of the bone, although in the close proximity to the suture between the basisphenoid and parietal. The relative shortness of the Vidian canals in one of the basisphenoids from Kohfidisch, along with its absolute size (Text-fig. 1: A-C), indicate that the bone belonged to an extremely large snake. On the dorsal side of each bone, paired foramina for the entry and exit of the abducens nerves are located around the crista sellaris in the manner characteristic of most snakes. The constrictor internus dorsalis branches of the trigeminal nerves (hereafter termed the cidnerves) entered the basisphenoid through narrow (eventually partially closed) grooves located between the lateral margins of the bone and the anterior orifices for the abducens nerves. The cid-nerves then re-entered the bone through foramina located on both sides of the pituitary fossa and left it cranial to the anterior orifices of the Vidian canals.

Two prootics from Kohfidisch come from the left and right sides of (a) braincase(s) (Text-fig. 2 A, B). The bone is subquadrate in shape; its roof possesses a strong crest for reception of the squamosal. The lateral wall contains two large cavities; the anterior for the maxillary branch of the trigeminal nerve (V2), and the posterior housing the mandibular branch of the nerve (V3) as well as the facial nerve; the cavities are accompanied by a number of smaller foramina. An orifice located below the V2 cavity admitted the cid-nerve to the basisphenoid. In the right bone the foramen is located in part within the V2 cavity. The facial nerve, leaving the V3 cavity by a smaller (i. e., more posterior) foramen, run towards the posterior orifice of the Vidian canal within a deep groove; in the left prootic the groove is partially closed by an extension of the laterospheroid portion of the bone.

The extension is pierced by the laterospheroid foramen.

©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at Neogene Cobras of the Genus Naja (Serpentes: Elapidae) of East Europe 57 Fig. 2.: Some bones oiNaja romani from Kohfidisch. A, right prootic in right lateral view (NMW 1989/ 34/4). B, left prootic in left lateral view (NMW 1989/34/5). C, basioccipital in posterolateral view (NMW 1989/34/3). D, parietal fragment in dorsal view (NMW 1989/34/3). E and F, left rib in ventral and inner views (NMW 1989/34/11). G, right quadrate in right lateral view (NMW 1989/34/9). H, left squamosal in left lateral view (NMW 1989/34/7). Scale line 5 mm. Abbreviations: V2, cavity for maxillary branch of trigeminal nerve; V3, cavity for mandibular branch of trigeminal nerve; V4, foramen for entry of cidnerve; VII, groove for facial nerve.

A basioccipital, a parietal fragment, two squamosals, two quadrates and numerous ribs from Kohfidisch (Text-fig. 2 C-H) are developed similarly to those of other cobras and do not possess important differentiating features.

Of the cranial elements belonging to the Kohfidisch fossil, described previously by BACHMAYER & SZYNDLAR (1985, 1987), the maxillae and a palatine are especially noteworthy. The venomous fangs on the maxillae are always followed by two solid teeth each. The palatine bears eight teeth, a very rare condition in Naja ©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at


(usually seven teeth). Several compound bones from Kohfidisch are very similar to those of other Naja. The single compound from Gritsev, despite its extensive damage, displays the same morphology.

Large vertebrae from Kohfidisch were described in detail by BACHMAYER & SZYNDLAR (1985). The vertebrae examined in the present study belonged to smaller (but adult) specimens (Text-fig. 3 A-C). The only differences between them are of allometric nature: the smaller vertebrae have relatively longer centra, wider neural canals and distinctly crenate zygosphenal roofs. The ten vertebrae from Gritsev (Text-fig. 3 D-E) do not differ significantly from those from Kohfidisch; they belonged to smaller specimens.

The vertebrae from Tourkobounia 1 (Text-fig. 3 G-I) display a set of different morphological features. They differ in having relatively broad centra, relatively Fig. 3.: Mid-trunk vertebrae of Naja romani from Kohfidisch (A-C; NMW 1989/34/12) and from Gritsev (D-F; IZAN 22-1776) and of Naja sp. from Tourkobounia 1 (G-I; UUGI uncat.); left lateral (A, D, G,), dorsal (B, E, H), and ventral (C, F, I) views. Scale line 5 mm.

©Naturhistorisches Museum Wien, download unter www.biologiezentrum.at Neogene Cobras of the Genus Naja (Serpentes: Elapidae) of East Europe 59 short and obtuse hypapophyses as well as convex (not crenate) zygosphenes in dorsal view. All vertebrae are damaged.

Discussion The vertebrae from Tourkobounia 1 undoubtedly belonged to a different species than those from Gritsev and Kohfidisch. Unfortunately, proper identification to the specific level cannot be demonstrated. In some details (especially the zygosphenal roof), they resemble vertebrae of the Asiatic N. naja but also those of some African forms, e. g., N. melanoleuca.

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