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Fossil Amphibians and Reptiles from the Neogene Locality of Maramena (Greece), the Most Diverse European Herpetofauna at the Miocene/Pliocene Transition Boundary

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Fossil Amphibians and Reptiles from the Neogene Locality of Maramena (Greece), the Most Diverse European Herpetofauna at the Miocene/Pliocene Transition Boundary Palaeontologia Electronica palaeo-electronica.org Fossil amphibians and reptiles from the Neogene locality of Maramena (Greece), the most diverse European herpetofauna at the Miocene/Pliocene transition boundary Georgios L. Georgalis, Andrea Villa, Martin Ivanov, Davit Vasilyan, and Massimo Delfino ABSTRACT We herein describe the fossil amphibians and reptiles from the Neogene (latest Miocene or earliest Pliocene; MN 13/14) locality of Maramena, in northern Greece. The herpetofauna is shown to be extremely diverse, comprising at least 30 different taxa. Amphibians include at least six urodelan (Cryptobranchidae indet., Salamandrina sp., Lissotriton sp. [Lissotriton vulgaris group], Lissotriton sp., Ommatotriton sp., and Sala- mandra sp.), and three anuran taxa (Latonia sp., Hyla sp., and Pelophylax sp.). Rep- tiles are much more speciose, being represented by two turtle (the geoemydid Mauremys aristotelica and a probable indeterminate testudinid), at least nine lizard (Agaminae indet., Lacertidae indet., ?Lacertidae indet., aff. Palaeocordylus sp., ?Scin- cidae indet., Anguis sp., five morphotypes of Ophisaurus, Pseudopus sp., and at least one species of Varanus), and 10 snake taxa (Scolecophidia indet., Periergophis micros gen. et sp. nov., Paraxenophis spanios gen. et sp. nov., Hierophis cf. hungaricus, another distinct “colubrine” morphotype, Natrix aff. rudabanyaensis, and another dis- tinct species of Natrix, Naja sp., cf. Micrurus sp., and a member of the “Oriental Vipers” complex). The autapomorphic features and bizarre vertebral morphology of Perier- gophis micros gen. et sp. nov. and Paraxenophis spanios gen. et sp. nov. render them readily distinguishable among fossil and extant snakes. Cryptobranchids, several of the amphibian genera, scincids, Anguis, Pseudopus, and Micrurus represent totally new fossil occurrences, not only for the Greek area, but for the whole southeastern Europe. The four different types of serration within the Varanus teeth from Maramena raise questions on the taxonomic importance or the variability of this feature. The large number of distinct amphibian and reptile taxa in Maramena makes this Greek locality by far the most diverse and speciose among all European localities across the latest Miocene and earliest Pliocene. An estimation of the palaeoprecipitation value of the locality is provided. The biogeographic origins of the Maramena herpetofauna are not fully resolved, though certain of its elements were previously only known from the early and middle Miocene of Central Europe. http://zoobank.org/5857474C-1265-4443-8938-BC37420561F1 Georgalis, Georgios L., Villa, Andrea, Ivanov, Martin, Vasilyan, Davit, and Delfino, Massimo. 2019. Fossil amphibians and reptiles from the Neogene locality of Maramena (Greece), the most diverse European herpetofauna at the Miocene/Pliocene transition boundary. Palaeontologia Electronica 22.3.68 1-99. https://doi.org/10.26879/908 palaeo-electronica.org/content/2019/2797-fossil-herpetofauna-maramena Copyright: November 2019 Society of Vertebrate Paleontology. This is an open access article distributed under the terms of the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original author and source are credited. creativecommons.org/licenses/by/4.0/creativecommons.org/licenses/by-nc-sa/4.0/ GEORGALIS ET AL.: FOSSIL HERPETOFAUNA MARAMENA Georgios L. Georgalis. Dipartimento di Scienze della Terra, Università di Torino, Via Valperga Caluso 35, 10125 Torino, Italy. [email protected] Department of Geosciences, University of Fribourg / Freiburg, Chemin du Musée 6, 1700 Fribourg, Switzerland. Department of Ecology, Laboratory of Evolutionary Biology, Faculty of Natural Sciences, Comenius University in Bratislava, Mlynská dolina, 84215 Bratislava, Slovakia. Andrea Villa. Bayerische Staatssammlung für Paläontologie und Geologie, Richard-Wagner-Straße 10, 80333 Munich, Germany. [email protected] Dipartimento di Scienze della Terra, Università di Torino, Via Valperga Caluso 35, 10125 Torino, Italy. Martin Ivanov. Department of Geological Sciences, Faculty of Science, Masaryk University, Kotlářská 2, 611 37 Brno, Czech Republic. [email protected] Davit Vasilyan. JURASSICA Museum, Fontenais 21, 2900 Porrentruy, Switzerland. Department of Geosciences, University of Fribourg / Freiburg, Chemin du Musée 6, 1700 Fribourg, Switzerland. [email protected] Massimo Delfino. Dipartimento di Scienze della Terra, Università di Torino, Via Valperga Caluso 35, 10125 Torino, Italy. [email protected] Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA-ICP, Carrer de les Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona, Spain. Keywords: amphibians; reptiles; Neogene; biogeography; new species; new genus Submission: 26 July 2018. Acceptance: 14 October 2019. INTRODUCTION ern portions of the continent (Szyndlar, 1991a, b, 2012; Ivanov et al., 2000; Rage, 2013). The latest Miocene/earliest Pliocene transition To fully comprehend the mechanisms around in Europe is characterized by important climatic these extinctions, probable dispersals, distribution changes, geographic alterations, and faunal turn- ranges alterations, and faunal turnovers of amphib- overs, which affected the palaeoenvironments of ian and reptile assemblages during the latest Mio- the continent (Cerling et al., 1997; Snel et al., cene/earliest Pliocene transition, it is essential to 2006; Carnevale et al., 2008; Harzhauser et al., comprehend and document in detail the exact 2015; Suc et al., 2015; Carnevale et al., 2017; diversity, taxonomic composition, and affinities of Karakitsios et al., 2017). One of the major dramatic palaeoherpetofaunas across European localities of events of the Cenozoic Era, the Messinian Salinity this time interval. Here we describe in detail the Crisis (hereafter MSC), took place at the latest Mio- fossil amphibians and reptiles from the locality of cene (Messinian), a period during which the Medi- Maramena, northern Greece. A large number of terranean Sea became isolated from the Atlantic amphibian and reptile lineages are described from Ocean through a closure of the Strait of Gibraltar, the fossil record of southeastern Europe for the first resulting in a significant amount of drying and time. Two new “colubrine” snake genera and spe- severe environmental perturbations (Hsü, 1972; cies are established to accommodate new taxa Krijgsman et al., 1999; Snel et al., 2006; Carnevale with bizarre autapomorphic features. With an age et al., 2008; Ivanovic et al., 2014; Harzhauser et around the boundary of latest Miocene/earliest al., 2015; Suc et al., 2015; Carnevale et al., 2017; Pliocene (MN 13/14), Maramena offers a unique Karakitsios et al., 2017; Carnevale et al., 2019). insight about the biogeography and taxonomic Although the main focus of palaeontological diversity of European herpetofaunal assemblages interest for this time lapse has been, as usual, fos- during that time interval. The high abundance and sil mammals (e.g., Koufos et al., 2005; Fortelius et diversity of taxa surpasses all contemporary fau- al., 2006; Koufos and Vasileiadou, 2015), studies nas of the continent and raises questions about the on fossil amphibians and reptiles have demon- negative effect of the Miocene/Pliocene transition strated a general impoverishement of post-Mio- on European herpetofaunas, implying a much cene European herpetofaunas, with certain clades more complex biogeographic pattern. being locally extinct from the continent and others gradually shrinking their ranges towards the south- 2 PALAEO-ELECTRONICA.ORG GEOLOGICAL AND PALAEOECOLOGICAL (Koufos, 2006). Three approximate and coeval SETTINGS sites in Maramena have produced fossil verte- brates, namely Maramena-1 (M1), Maramena-2 The locality of Maramena is situated a few (M2), and Maramena-3 (M3). The majority of the kilometers from the city of Serres, in the adminis- herpetofaunal material described herein, as also trative region of Central Macedonia, in northern the most complete specimens, originates from the Greece (Figure 1). Maramena, along with the adja- M1 site. cent localities Ano Metochi-2 and Ano Metochi-3, Maramena is so far mostly known for its abun- belongs to the Lefkon Formation (Bruijn, 1989). dant and diverse micromammal fauna, comprising Maramena is situated in the uppermost part of the erinaceomorphs, soricomorphs, lagomorphs, and Lefkon Formation, which consists of arkosic sands, numerous rodents, though large mammals have clays, marls, and lignites (Schmidt-Kittler et al., also been found, consisting of primates, carniv- 1995). The geology of the Lefkon Formation and orans, proboscideans, perissodactyls, and artio- the adjacent ones (Georgios Formation and Spilia dactyls (Bruijn, 1995; Schmidt-Kittler et al., 1995; Formation) within the Strymon (or Strimon or Koufos, 2006). No fossils of amphibians have been Serres) Basin is described in detail by Karystineos so far formally described from Maramena. As far as (1984), Bruijn (1989), and Schmidt-Kittler et al. it concerns reptiles, previous studies conducted in (1995). Further information regarding the Strymon the past three decades have documented the pres- Basin has been recently provided in detail by ence of fragmentary remains of turtles (Gad, Karakitsios et al. (2017). Maramena is considered 1990), lizards (Richter, 1995), and snakes (Szynd- to pertain either to the latest Miocene (latest Mes- lar, 1995). sinian)
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