Estudios geol., 47: 237-266 (1991) A REVIEW OF NEOGENE AND QUATERNARY SNAKES OF CENTRAL AND EASTERN EUROPE. PART 11: NATRICINAE, ELAPIDAE, VIPERIDAE Z. Szyndlar ABSTRACT Remains of Neogene and Quaternary "natricine" colubrids, elapids and viperids, inc­ luding snakes previously described and those undescribed yet, coming from Poland, Ukrai­ ne, Moldavia, Czechoslovakia, Austria, Hungary, Romania, Bulgaria, and Greece are dis­ cussed. The following taxa, including 11 extinct species, were recognized: "Natricinae": Neonatrix nova, Neonatrix sp., Palaeonatrix silesiaca, Palaeonatrix lehmani, Natrix longiv­ ertebrata, Natrix cf. N. longivertebrata, Natrix natrix, Natrix tesselata, Natrix cf. N. tessela­ ta, Natrix sp., "Natricinae" indet.; Elapidae: Naja romani, Naja sp., cf. Naja sp.; Viperi­ dae: Vipera platyspondyla, Vipera sarmatica, Vipera burgenlandica, Vipera gedulyi, Vipera kuchurganica, Vipera antiqua, Vipera cf. V. ammodytes, Vipera berus, Vipera sp. ('Oriental vipers' group), Vipera sp. ('aspis' group), Vipera sp. ('berus' group), Vipera sp. (status un­ known). Taxonomic status of two other extinct species, Natrix parva and Laophis crotaloi­ des, is uncertain. Modern species appeared fírst in Central and East Europe in the middle Pliocene (MN 15). Older snakes belonged to extinct species of either extinct or extant ge­ nera; taxonomic distinction of most extinct genera is, however, not fully demonstrated. Best recognized oldest snakes from the area (Elapidae, Viperidae, and sorne Colubridae) are clearly referable to modern genera and intrageneric subdivisions occurring today are ob­ served in oldest (Iower Miocene) remains; closest living relatives of these fossils are pre­ sently distributed in the Oriental Realm. Key words: Serpentes, Colubridae, Elapidae, Viperidae, Europe, Miocene, Pliocene, Pleis­ tocene, Taxonomy, Paleofaunistics, Morphology. RESUMEN Se revisan y estudian los restos neógenos y cuaternarios de colúbridos «natricinos», elá­ pidos y vipéridos, incluyendo tanto serpientes previamente descritas como- otras inéditas. Los materiales analizados proceden de Polonia, Ukrania, Moldavia, Checoslovaquia, Aus­ tria, Hungría, Rumania, Bulgaria y Grecia. Se reconocen los siguientes taxones, incluyen­ do 11 especies extinguidas: Natricinae: Neonatrix nova, Neonatrix sp., Palaeonatrix silesia­ ca, Palaeonatrix lehmani, Natrix longivertebrata, Natrix cf. N. longivertebrata, Natrix na­ trix, Natrix tesselata, Natrix cf. N. tesselata, Natrix sp., «Natricinae» indet.; Elapidae: Naja romani, Naja sp., cf. Naja sp.; Viperidae: Vipera plastyspondyla, Vipera sarmatica, Vipera burgenlandica, Vipera gedulyi, Vipera kuchurganica, Vipera antiqua, Vipera cf. V. ammody­ tes, Vipera berus, Vipera sp. (<<grupo Oriental»), Vipera sp. (grupo «aspis»), Vipera sp. (gru­ po «berus»), Vipera sp. (relaciones desconocidas). El estatus taxonómico de otras dos es­ pecies extintas, Natrix parva y Laophis crotaloides, es poco claro. Las especies actuales apa­ recen primero en Europa central y oriental durante el Plioceno medio (MN 15), mientras que las serpientes más antiguas pertenecen a especies extinguidas, incluidas tanto en géne­ ros actuales como también en otros ya extinguidos. La validez taxonómica como entes in­ dependientes de muchos géneros extintos no está plenamente demostrada. Las serpientes bien conocidas más antiguas del área (Elapidae, Viperidae, algunos Colubridae) son clara­ mente adscribibles a géneros actuales, y las subdivisiones intragenéricas actualmente exis­ tentes pueden ya observarse en los restos más antiguos (Mioceno inferior). Formas actua­ les cercanas a estos fósiles miocénicos se encuentran distribuidos en la Region Biogeográ­ fíca Oriental. Palabras clave: Serpentes, Colubridae, Elapidae, Viperidae, Europa, Mioceno, Plioceno, Pleistoceno, Taxonomía, Paleofaunística, Morfología. * Polish Acaderny of Sciences, Institute of Systernatics and Evolution of Anirnals, Slawkowska 17, 31-016 Kraków. Poland. 238 Z. SZYNDLAR Introduction from those belonging to other hypapophysis-bearing snakes. They differ from the Viperidae in having usuaIly sigmoid (and not straight) hypapohyses, posteriorly vaulted (and not depressed) Toe present paper, devoted to Neogene and Qua­ neural arches, shorter parapophyseal processes, and usuaIly much ternary "natricine" colubrids, elapids and viperids longer centra (Szyndlar 1984, 1988). They differ from the Elapi­ from Central and Eastern Europe, supplements dae in having lightly built vertebrae, provided with much longer centra and strong subcentral ridges. In addition, the modero ge­ Szyndlar's (1991) review of fossil scolecophidians, nus Natrix has relatively much higher neural spines than those of boids and "colubrine" colubrids coming from the elapids. same area. Table 1 lists aH ophidian taxa described Natricine snakes presently inhabiting Europe belong exclusive­ Iy to the genus Natrix (see below). There are two valid extinct ge­ in both papers. nera known from Europe, Neonatrix and Palaeonatrix. The arrangement of the information presented be­ low in the chapter "Systematic account" foHows Genus Neonatrix Holman, 1973. exactly the system employed in my previous paper; Neonatrix is an extinct genus, widespread in the North Ameri­ see chapters "Localities" and "Systematic account" can and European Miocene (Holman, 1979; Rage, 1984; Rage and Holman, 1984). The most important diagnostic feature of this ge­ in Szyndlar (1991) for details. nus are strongly reduced hypapophyses. One species was reported The chapter "History of snakes in Central and East from East Europe. Europe" is based on the data presented in both pa­ pers and summarizes information about the compo­ Neonatrix nova Szyndlar, 1987 (fig. 1). sition and past distribution of the extinct snake fau­ 1987a Neonatrix nova Szyndlar, pp. 61-62, fig. 6. na in the area, against the background of the recent Material: (37) Lower Miocene (MN 4) of Dolnice (type loca­ lity): one trunk vertebra (holotype, DPFNSP 5197), 62 precaudal fauna. The last chapter of the present paper, contain­ vertebrae (DPFNSP 931, 932, 934-937, 1171, 1237, 1245, 1253, ing a critica! review of methods used in ophidian pa­ 1311, 1312, 1414, 1419, 1430, 1433, 1440, 1443, 1451, 1458, 3962, leontology, summarizes my ten-years experience with 3967,3979,3983,3987,3988,4026,4029,4030,4075,4107,4201, research on Neogene and Quaternary European sna­ 4511, 4526, 4532, 5156-5159, 5162, 5164, 5169, 5171-5173, 5175, 5176, 5182, 5186-5188, 5192, 5194-5196, 5198, 5200, 5202, 5204, kes. 5214, 5221, 5224). Diagnostic vertebral characters: This extinct species, known only from the type locality, is most similar to its North American relative, N. elongata, but differs from the latter in much larger ab­ Systematic Account solute size. The centrum length of the holotype vertebra is 4.33 mm and it is 1.54 times longer than wide. For detailed des­ Family COLUBRIDAE (s.l.) Oppel, 1811 cription see Szyndlar (1987a). "Natricines". The basic feature differentiating natricine vertebrae from those Neonatrix sp. of other members of the family Colubridae (s.l.) is the presence Material: (38) Middle Miocene (MN 6) of Devínska Nová Ves: of hypapohyses throughout the precaudal region of the column; 3 precaudal vertebrae (DPFNSP 5835,5837,5843). colubrines possess hypapophyses in the cervical region only, while Remarks: The vertebrae, somewhat damaged, c10sely resemble in the remaining precaudal vertebrae this structure is replaced by those of N. nova, but differ from them by smaIler absolute size the haemal keel. Natricine vertebrae can be easily distinguished and by having strongly reduced neural spines. ] Fig. I.-Trunk vertebra of Neonatrix nova from lower Miocene of Dolnice (holotype, DPFNSP 5197). A, left lateral view; B, dor­ sal view; C, ventral view; D, anterior view; E, posterior view. Sca­ le equals 2 mm (From Szyndlar, 1987a. Copyright 1987 by the So- ciety of Vertebrate Paleontology. Used with permission). A REVIEW OF NEOGENE AND QUATERNARY SNAKES OF CENTRAL AND EASTERN EUROPE 239 Genus Palaeonatrix Szyndlar, 1982. dition, four fossil species are known from Europe; their geologi­ cal age ranges from the Oligocene to Upper Pliocene (Rage 1988). This extinct genus consists of two species, P. silesiaea and P. leh­ The most important feature differentiating Natrix from the extinct mani, both from the East European Miocene. These snakes are natricine genera is a distinctly higher neural spine. known exclusively by vertebrae. The most important diagnostic features, differentiating Palaeonatrix from the living genus Natrix Natrix longivertebrata Szyndlar, 1984 (fig. 3). and sorne other related genera, are a prominent anterior keel ac­ companying the hypapophysis and a vestigial neural spine (6) 1984 Natrix longivertebrata Szyndlar, pp. 71-80, figs. 27, (Szyndlar 1987a). 29. (34) 1985 Natrix longivertebrata Szyndl.: David et al., p. 73. Palaeonatrix silesiaea Szyndlar, 1982. (6) 1986 Natrix longivertebrata: Rage and Szyndlar,passim, figs. 2, 4, 7, 8, 11, 12, 15, 16. 1982 Palaeonatrix silesiaea Szyndlar (in Mlynarski et al.), pp. (31) (32) 1987 Natrix longivertebrata: Redkozubov, p. 71. 114-116, fig. 10. (34) 1989 Natrix longivertebrata: Redkozubov, p. 209. 1984 Palaeonatrix silesiaea Szyndlar: Szyndlar, pp. 45-47, fig. 16. Material: (31) Middle Pliocene (MN 15) of Musait: trunk ver- Material: (2) Middle Miocene (MN 7) of Opole 2 (type loca­ tebrae (not seen, fide Redkozubov 1987). (6) Upper Pliocene lity): one trunk vertebra (holotype, ZPUW OP-86/24), 5,erecau­ (MN 16) of Rc;bielice