First Description of a Fossil Chamaeleonid from Greece and Its Relevance for the European Biogeographic History of the Group

Published in 7KH6FLHQFHRI1DWXUH ± ± which should be cited to refer to this work. First description of a fossil chamaeleonid from Greece and its relevance for the European biogeographic history of the group Georgios L. Georgalis1,2 & Andrea Villa2 & Massimo Delfino2,3 Abstract The fossil record of Chamaeleonidae is very scarce Introduction and any new specimen is therefore considered important for our understanding of the evolutionary and biogeographic his- Chameleons (Squamata, Chamaeleonidae) constitute a tory of the group. New specimens from the early Miocene of rather diverse group with more than 200 species that are Aliveri (Evia Island), Greece constitute the only fossils of now distributed in Africa, Madagascar and several other these lizards from southeastern Europe. Skull roofing material Indian Ocean islands, southern Asia, Cyprus and southern is tentatively attributed to the Czech species Chamaeleo cf. parts of Mediterranean Europe (Glaw 2015). Cryptic di- andrusovi, revealing a range extension for this taxon, whereas versity appears to be prominent within the group, with tooth-bearing elements are described as indeterminate several new species having been described only during chamaeleonids. The Aliveri fossils rank well among the oldest the current decade, mostly on the basis of molecular data known reptiles from Greece, provide evidence for the dispers- (Gehring et al. 2011; Crottini et al. 2012; Greenbaum al routes of chameleons out of Africa towards the European et al. 2012;Stipalaetal.2011, 2012; Tilbury and Tolley continent and, additionally, imply strong affinities with coeval 2015). The size range of chamaeleonids is also broad and chamaeleonids from Central Europe. astonishing, with the larger members of the family sur- passing 600 mm in total length (Glaw and Vences 1994), whereas the smallest species rank well among the Keywords Chamaeleonidae . Squamata . Miocene . tiniest known reptiles, both extant (Brookesia micra, . http://doc.rero.ch Biogeography Aliveri attaining only 29 mm [Glaw et al. 2012]) and extinct (e.g. Jucaraseps grandipes [Bolet and Evans 2012]). The ability of some species to ‘change’ colour, along with Electronic supplementary material The online version of this article their peculiar external morphology, have gained the atten- contains supplementary material tion of the public for centuries and render them readily distinguishable even for non-specialists (Teyssier et al. 2015). Moreover, their bizarre external appearance, * Georgios L. Georgalis acrodont dentition, unique skeletal anatomy, as even their [email protected] specialized feeding and locomotor habits have long lead herpetologists to consider chameleons as a monophyletic 1 group (Camp 1923; Estes 1983). Their relationships to Department of Geosciences, University of Fribourg/Freiburg, other squamates, however, are yet not fully resolved, Chemin du Musée 6, 1700 Fribourg, Switzerland 2 mostly because of the uncertainty of the position of Dipartimento di Scienze della Terra, Università di Torino, Via Iguania, but it is widely accepted that Chamaeleonidae Valperga Caluso 35, 10125 Torino, Italy 3 constitute a group within the iguanian clade Acrodonta Institut Català de Paleontologia Miquel Crusafont, Universitat (Estes et al. 1988;Townsendetal.2004; Kumazawa Autònoma de Barcelona, Edifici ICTA-ICP, Carrer de les Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, 2007; Conrad 2008; Vidal and Hedges 2009; Gauthier Barcelona, Spain et al. 2012; Wiens et al. 2012; Pyron et al. 2013; Reeder 1 et al. 2015), even if their affinities with Agamidae are a MN 3 zone (de Bruijn et al. 1980). Nevertheless, there matter of debate, with different topologies arising on the is a general consensus now that it is referable to MN 4 basis of morphological and molecular data (Bolet and (Doukas 2003;Koufos2006b; van den Hoek Ostende Evans 2013; Tolley and Menegon 2013). et al. 2015). More specifically, as was recently stated by Despite the broad geographic distribution among extant van den Hoek Ostende et al. (2015:276), ‘although unde- taxa, the fossil record of Chamaeleonidae is rather patchy niably an MN 4 assemblage, Aliveri is one of the oldest (Bolet and Evans 2013). A Cretaceous origin of the group is localities from that MN unit, and presumably co-eval with commonly accepted based on molecular data (Tolley et al. many of the MN 3 assemblages from western and central 2013), but with their first fossil record dating back to the early Europe’. The age of Aliveri is tentatively estimated be- Miocene (Moody and Roček 1980; Pickford 1986), such mo- tween 18 and 17.5 Ma, and corresponds to the lecular divergence date cannot be thoroughly tested. The dis- Burdigalian stage (Orleanian Continental Stage) (Koufos persal of chamaeleonids to Europe and Asia seems to have 2006b; van den Hoek Ostende et al. 2015). A detailed occurred later (Bolet and Evans 2013; Tolley et al. 2013); description of the geology of Aliveri is provided by de indeed, their Neogene record on the European continent is Bruijn et al. (1980). rather scanty with few, scattered occurrences only across the The micromammal assemblage of Aliveri is rather di- early to middle Miocene of Central Europe (Moody and verse and has been the subject of several studies (de Roček 1980; Böhme and Ilg 2003; Čerňanský 2010;Bolet Bruijn et al. 1980; van der Meulen and de Bruijn 1982; and Evans 2013). Schmidt-Kittler 1983; Klein Hofmeijer and de Bruijn Europe currently harbours two extant taxa of 1985, 1988; Doukas 1986; López Martinez 1986; chamaeleonids: Chamaeleo chamaeleon (Linnaeus, 1758) Álvarez Sierra et al. 1987). It includes erinaceids, and Chamaeleo africanus Laurenti, 1768 (Dimaki et al. dimylids, heterosoricids, talpids, soricids, ochotonids, 2000, 2008, 2015). The Greek distribution of the former spe- eomyids, many sciurids, cricetids, anomalomyids, cies now solely comprises Samos island, as populations spalacids and glirids. Larger mammals are less known, inhabiting Crete and Chios seem to have gone extinct but palaeogalids, equids, palaeomerycids, cervids and bo- recently. It is worth noting, however, that Sillero et al. vids are present, as well as the viverrid Euboictis (van den (2014) reported this species as still present in Crete, consider- Hoek Ostende et al. 2015). As for the herpetofauna of ing it allochthonous. The distribution of C. africanus in Aliveri, anurans, crocodylians, anguids, lacertids, as well Greece (restricted only to a small region near the town of as indeterminate lizards, snakes and turtles are present but Pylos, Peloponnese) is the sole European population for this have not been described. The study of these amphibians African species and most probably is a product of human and reptiles will be the subject of a forthcoming introduction during early antiquity (Fig. 2)(Böhmeetal. publication. 1998;Kosuchetal.1999;Dimakietal.2000, 2008). Despite their occurrence in the extant herpetofauna of the country, chamaeleonids have never been previously described Material and methods http://doc.rero.ch from Greek fossil localities. Here, we report on three new chamaeleonid fossils found in the early Miocene (MN 4) lo- All three chamaeleonid specimens described herein belong to cality of Aliveri, Evia Island, which indicate the Neogene the collections of the Department of Earth Sciences in the existence of these lizards in Greece. One of these specimens University of Utrecht and are accessioned under the repository is tentatively attributed to the species Chamaeleo cf. numbers UU AL 3501–3503. All specimens were found in the andrusovi Čerňanský, 2010, which was previously known early Miocene locality of Aliveri in three different field sea- only from the early Miocene of the Czech Republic, whereas sons, in 1977 (UU AL 3502), 1978 (UU AL 3501), and 1980 the other two specimens represent indeterminate (UU AL 3503), led by the University of Utrecht. As with the chamaeleonids. micromammal material that has also been collected from this locality, all chamaeleonid fossils have been recovered from Geological setting the underclay of the lignite occurring North of Aliveri, along the eastern edge of the basin where the lignite wedges out Aliveri is a fossiliferous locality, located within the against Mesozoic limestones (de Bruijn et al. 1980). For com- Neogene sedimentary basin of Aliveri/Kymi, in Evia (also parison purposes, skeletal material of extant chamaeleonids spelled Euboea) Island in Central Greece. Along with the was studied in the herpetological collections of the Gavathas site, Lesvos Island, it has yielded the oldest University of Torino and the Muséum national d’Histoire known Greek Neogene land mammal fauna (Koufos naturelle, Paris. Photos of the fossil specimens were taken 2006a, b; van den Hoek Ostende et al. 2015). Aliveri using the Leica M205 microscope and the Leica application was discovered in 1977 and was initially dated to the suite V 3.3.0, in the University of Torino. 2 Results the four clusters are aligned along one of the margins of the fragment, which is a natural border and not the result of break- Systematic palaeontology age (as the other margins). If our interpretation of the fragment as part of a circumorbital bone is correct, this would be the Reptilia Laurenti 1768 orbital margin. The inner surface of the specimen is smooth. Squamata Oppel, 1811 Chamaeleonidae indet. Iguania Cope, 1864 (Fig. 1e–h) Acrodonta Cope, 1864 Material: Two additional specimens from the same locality Chamaeleonidae Gray, 1825 are assigned to Chamaeleonidae, but cannot be attributed to Chamaeleo Laurenti, 1768 the species level. These two specimens are both tooth bearing Chamaeleo cf. andrusovi Čerňanský, 2010 bones (UU AL 3502 and UU AL 3503). (Fig. 1a, b, c, d) Description: UU AL 3502 and UU AL 3503 are very frag- Material: The specimen here referred to Chamaeleo cf. mentary tooth-bearing bones. However, they clearly bear andrusovi is a skull roofing bone fragment (UU AL 3501). acrodont, large and triangular teeth, which are well-spaced Description: UU AL 3501 is a skull roofing bone fragment and located on the edge of the bones. Two of them are pre- about 3 × 3 mm in diameter.

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