Khosrovsky reservation, and in Turkmenistan on the mountain Dushack in the central Kopetdag, blunt-nosed vipers are found at the altitude up to 2000 m above sea level. On the Pamir, at the altitude 2500 m above sea level, populations from even higher mountains are known (Ananjeva et al. 2006). The Armenian viper, which can be over 1 m in length, occurs at the altitude 1000–2700 m above sea level in mountain-xerophytic vegetation, in particular oak forests, in the juniper open woodlands, on the rocky slopes of the mountains with sparse bush vegetation, and in the mountain steppes. The type of biotope distribution on the southern spurs of the mountain ranges of the Lesser Caucasus corresponds to the mosaic availability of suitable biotopes (Ananjeva et al. 2006).

In the Miocene, “Oriental vipers” were widely distributed in the entire southern half of Europe and survived until the end of the Pliocene in the Mediterranean area and in Eastern Europe until at least the Middle Pleistocene (Szyndlar 1991b). In Asia Minor Vipera lebetina is known in the Middle to Late Pleistocene of Karain E, Turkey (S. Bailon, personal communication) and in the Late Pleistocene from Wezmeh Cave, Iran (Mashkour et al. 2009). An “Oriental viper” (Vipera sp.) has been mentioned too in the Middle Pleistocene of Emirkaya-2, Turkey (Venczel and Sen 1994).

Paleobiogeographical Data

Some Palearctic regions were particularly important as corridors for invasions. Trans-Caucasia was repeatedly affected by fauna and flora exchanges because it is situated at the interface of the European, Asian and African biomes from where it experienced repeated invasions since the Late Oligocene (Veith et al. 2003). According to Sindaco et al. (2000), this area is currently characterized by a high number of endemic reptile taxa bearing an “Armenian” pattern of distribution, which usually includes species inhabiting mountains or plateaux and usually adapted to steppe or rocky habitats. Excluding these endemic species, three chorotypes are dominant: the Southwestern-Asiatic, the Eastern-Mediterranean and the Turano-Mediterranean.

According to ecologists and biologists (Ministry of Nature Protection 1999), a series of climatic modifications have played an important role in determining the current landscapes, ecosystems and biodiversity of the Caucasus region. During the Late Pliocene and Pleistocene, a series of glaciations occurred which affected the existing ecosystems, resulting in reductions in primary subtropical forests, and growth in secondary habitats typical of the temperate zone. During cold phases, reptiles of European origin, such as the meadow viper (Vipera ursinii), grass snake (Natrix natrix), sand lizard (Lacerta agilis), and meadow lizard

(L. praticola) migrated into the Armenian Plateau. During temperate or warm phases, the flora and fauna became dominated by taxa adapted to xerophilous conditions (probably with an Eastern-Mediterranean origin), as more arid-zone habitats emerged. A number of reptiles may also have reached the Armenian Plateau from the Middle Asian deserts, including the race runner (Eremias arguta), pond turtle (Mauremys caspica), and toadhead agama (Phrynocephalus persicus).

Most of the taxa represented in Azokh 1 have clear Eastern-Mediterranean or Turano-Mediterranean affinities (like P. syriacus, P. apodus, O. elegans, E. sauromates), whereas some others have a wider distribution area but always including in the Turano-Mediterranean area (P. viridis and E. jaculus). As an exception C. austriaca and the representatives of the V. berus complex have a more European or Sibero-European distribution and may constitute a special case for the Caucasian area which served as a refuge during the Pleistocene coldest periods.

European and Turano-Mediterranean species have been present at least since the Middle Pleistocene (Unit III). Only Mediterranean amphibians and reptiles are represented in the Early Pleistocene of Dmanisi, Georgia (Delfino in Lordkipanidze et al. 2007) and some Sibero-European taxon (Lacerta cf. agilis and Natrix cf. natrix) have been mentioned during or before the Pliocene in the South of the Russian Platform (Ratnikov 2009). On the contrary no clear Middle Asian desert taxa have been found in Azokh 1, but this may be partly the result of the lack of evidence for the systematic position of the small lacertid and agamid lizards, which does not allow us to see if any of the lizards had middle Asian affinities. Nevertheless, it is noteworthy that they only appear in Unit I (i.e. very recently), which may be due to the fact that more arid-zone habitats only emerged after glaciation.

Paleoclimatological

and Paleoenvironmental Inferences

Before completing a quantitative reconstruction of the environment, the origin of the small vertebrate accumulation (i.e. taphonomy) must be understood. During the systematic descriptions, strong evidence of digestion has been seen, in particular on some trunk vertebrae of erycine snake, suggesting predation by a small carnivore. However some preliminary remarks can be made.

The herpetofauna of Azokh 1 is composed exclusively of extant genera and species, the majority of them belonging to thermophilous and xerophilous forms (e.g., Pelobates syriacus, Agamidae, Pseudopus apodus, Ophisops elegans, Eryx jaculus, Elaphe sauromates, etc.). The anuran Pseudepidalea

viridis is a form with a wide ecological tolerance. Many of the taxa of Azokh 1 may frequent open wooded or bushy areas. Moreover high mountainous taxa are well represented, with the presence of a representative of the V. berus complex (probably V. ursinii) and the small colubrine Coronella austriaca. There are two high altitude mountains (around 2000–2500 m) within 6–10 km of Azokh, but the site itself is at less than 1000 m and the surrounding mountains barely reach 1500 m (P. Andrews 2010, personal communication). Through the sequence, the faunas from Units I to II appear to represent the driest period with the exclusive presence of agamid lizards and P. syriacus, whereas the unit Vu fauna seems to indicate “moister” conditions due to the presence of a larger number of C. austriaca remains. Moreover in the Unit Vu fauna, there are no species present that avoid forest environments, such as P. syriacus and the small vipers. So the environment in Unit Vu seems to be more consistent with a meadow-steppe whereas in Units III, II and particularly Unit I, the environment seems to have been more xeric corresponding to an arid mountain steppe (now occurring at lower elevations than meadow-steppes). Here is a case that demonstrates that need to understand the taphonomy, for if the herpetofauna represents mountain steppe, it must have been transported from considerable distances and it needs to be shown how it reached the cave. The small mammal fauna shows a similar range of ecological adaptations, but it has been shown to have been transported to the cave by predatory birds, which can easily travel these distances (Andrews et al. 2016). The climate seems to have been always relatively warm-temperate.

Conclusions

1.The amphibian and squamate reptile fossil fauna from the 2002–2009 excavations of Azokh 1 is composed of 14 taxa made up of three anurans (Pelobates cf. syriacus, Pseudepidalea viridis and Ranidae/Hylidae indet.), at least fave lizards (Agamidae indet., Pseudopus apodus, Lacerta sp., Ophisops elegans and Lacertidae indet.) and seven snakes [Eryx jaculus, cf. Coronella austriaca, cf. Elaphe sp. (probably E. sauromates), cf. “Coluber” sp., “Colubrinae” indet., Vipera sp. [V. berus complex (probably V. ursinii)], Vipera sp. (“Oriental vipers” complex or

Daboia)].

2.The herpetofauna of Azokh 1 cave is composed exclusively of extant genera and species, the majority of them belonging to thermophilous and xericadapted forms.

3.Most of the taxa have clear Eastern-Mediterranean or Turano-Mediterranean affinities, whereas some others have a larger distribution area but always including in the Turano-Mediterranean region.

4.A notable exception is the presence of C. austriaca and the representatives of the V. berus complex that have European or Sibero-European affinities. Azokh 1 represents the first fossil evidence for their presence in the Caucasian area at around 200 ka (Units III and V-upper).

5.From a chronological point of view, European and Turano-Mediterranean species are present at least since the Middle Pleistocene (Unit Vu) whereas no clear Middle Asian desert taxon has been found in Pleistocene levels at Azokh 1 (since the small lacertids and agamid lizard systematic affinities have yet to be elucidated). They only appear in Holocene deposits of Unit I (i.e. very recently), and this may be due to the fact that more arid-zone habitats emerged in the Armenian Plateau after the last glaciation.

6.From a paleoenvironmental point of view, the environment at the time of Unit Vu is consistent with a meadow-steppe environment, whereas in Unit III, II and particularly in Unit I, the environment may have been more xeric corresponding to an arid mountain steppe (now occurring at lower elevations that meadow-steppe). The climate seems always to have been relatively warm-temperate.

Acknowledgements We are deeply grateful to Dr. Prof. Vyacheslav Yu. Ratnikov (State University of Voronezh, Russia), Dr. Prof. Zbigniew Szyndlar (Institute of Systematics and Evolution ofAnimals of the Polish Academy of Sciences, Krakow, Poland), Dr. Salvador Bailon (MNHN, Paris, France), Dr. Massimo Delfino (University of Turin, Italy), Dr. Jean-Claude Rage (MNHN, Paris, France), Dr. Marton Venczel (Museul Tárii Crisurilor, Oradea, Romania), Dr. Borja Sanchiz (MNCN, Madrid, Spain), Dr. Yolanda Fernández-Jalvo (MNCN, Madrid, Spain) and to Dr. Peter Andrews (The Natural History Museum, London, UK) for providing us bibliography and/or unpublished information. Dr. José Enrique González and Dr. Salvador Bailon permitted us to consult the osteological material for comparison respectively from the Amphibians and Reptiles collections of the

Museo Nacional de Ciencias Naturales in Madrid and from the Anatomie Comparée collections of the Muséum national d’Histoire naturelle in Paris. We want to thank Dr. Prof. Paloma Sevilla (Universidad Complutense de Madrid, Spain), Lena Asryan and Sandra Bañuls (IPHES, Tarragona, Spain) for continuous encouragement and pleasant discussions on Azokh localities and Armenia. Dr. Marton Venczel, Dr. Prof. Zbigniew Szyndlar and Dr. Salvador Bailon as well as two of the editors Dr. Yolanda Fernández-Jalvo and Dr. Peter Andrews improved this manuscript by their comments.

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