The antiquity of Asian chameleons—first potential Chamaeleonidae and associated squamate fauna from the Lower and Middle Miocene of Anatolia

ABSTRACT

We here describe fossil remains from three Lower and Middle Miocene localities of Anatolia, which we tentatively refer to chamaeleonids. The material comprises isolated tooth-bearing bones from the Early Miocene (MN 3) of Sabuncubeli (western Anatolia), the Early Miocene (MN 4) of Dededağ (western Anatolia), and the Middle Miocene (MN 5) of the world-renowned hominoid locality of Paşalar (northwestern Anatolia). If our identifications are correct, these specimens would represent the only fossil record of this group from Anatolia as well as the oldest occurrences of chameleons from Asia. Moreover, the recovery of these specimens across different Lower and Middle Miocene localities, suggest that these animals were a typical element of the Anatolian herpetofaunas of the early Neogene. Differences among the specimens from the three localities point to a higher taxonomic diversity of Miocene Anatolian chamaeleonids, however, based on such limited material, intraspecific or ontogenetic variation cannot be ruled out. The oldest among these occurrences, the Sabuncubeli material, coincides chronologically with the oldest chameleons, which had been previously described from Central Europe (MN 3). As such, this supports the recently suggested biogeographic scenario, according to which chamaeleonids dispersed from Afro-Arabia to Europe during the Burdigalian, via the “Gomphotherium Landbridge,” through Anatolia and the Balkans. Finally, a few additional lizard and snake remains from the locality of Dededağ are described, adding to the poorly known fossil herpetofaunas of Anatolia.

ACKNOWLEDGMENTS

GLG acknowledges funding from the Ulam Program of the Polish National Agency for Academic Exchange (PPN/ULM/2020/1/00022/U/00001) and the Forschungskredit of the University of Zurich, grant no. [FK-20-110]. AČ acknowledges funding from the Scientific Grant Agency of the Ministry of Education of Slovak Republic and Slovak Academy of Sciences, Grant Nr. 1/0191/21. SM was supported by Ege University Research Projects, TTM/001/2016 and TTM/002/2016. The Paşalar excavations were carried out on behalf of the T.C. Ministry of Culture and Tourism, Directorate of Antiquities, and the University of Ankara, Faculty of Language, History, and Geography. Excavations were funded by the Directorate of Antiquities. GLG, AŞ, and SM particularly thank B. Alpagut for inviting us to work with her on the Paşalar chameleon material. For access to further comparative skeletal material of extant taxa, we would like to thank M. Delfino (MDHC), B. Borczyk (MNHW), and J. Moravec (NMP). The map of Fig. 1 was adapted from Wikimedia Commons (https://commons.wikimedia.org/wiki/File:Europe_blank_laea_location_map.svg; License CC-BY-SA-3.0). The quality of the paper was enhanced by the useful comments and suggestions provided by the Editor E. Maxwell and two anonymous reviewers.

AVAILABILITY OF MATERIALS AND DATA

All fossil specimens described herein are permanently curated at the collections of EUNMH PV and BP. The 3D models of all the extant specimens (except for that of Draco volans, PRIF UK AČ 08) are deposited at the Morphosource repository (https://www.morphosource.org/): right mandible (PRIF UK (AČ) DE 74) of Chamaeleo calyptratus (Media 000482071; ark:/87602/m4/482071); right mandible (PRIF UK AČ 82) of Calumma globifer (Media 000481987; ark:/87602/m4/481987); right mandible (PRIF UK AČ 01) of Trioceros jacksonii (Media 000482065; ark:/87602/m4/482065); right mandible (PRIF UK AČ 04) of Trioceros melleri (Media 000481981; ark:/87602/m4/481981); right mandible (PRIF UK AČ 80) of Furcifer pardalis (Media 000481993; ark:/87602/m4/481993); right mandible (PRIF UK AČ 06) of Bronchocela jubata (Media 000481974; ark:/87602/m4/481974); right mandible (PRIF UK AČ 07) of Uromastyx sp. (Media 000481968; ark:/87602/m4/481968).