The oldest known European Neogene girdled lizard fauna (Squamata, Cordylidae), with comments on Early Miocene immigration of African taxa Andrej ČERŇANSKÝ Senckenberg Research Institute and Natural History Museum Frankfurt, Palaeoanthropology and Messel Research, Senckenberganlage 25, D-60325 Frankfurt am Main (Germany) and Geological Institute, Slovak Academy of Sciences, Dumbierska ̌ 1, 974 01 Banská Bystrica (Slovak Republic) [email protected] Čerňanský A. 2012. — The oldest known European Neogene girdled lizard fauna (Squamata, Cordylidae), with comments on Early Miocene immigration of African taxa. Geodiversitas 34 (4): 837-848. http://dx.doi.org/10.5252/g2012n4a6 ABSTRACT This paper reports on the first record of cordylid lizards from the locality of Merkur-North. The fossil history of girdled lizards is very poorly known and this group was rare during the Lower Miocene in Europe. The fossils described herein come from grey calcareous marls at the base of the so-called “Main Brown Coal Seam”. These marls are interpreted as reworked volcanic ash and the sedi- ments are considered as early Miocene in age and are precisely equated with the MN 3a zone. For this reason, the cordylid fauna from this locality is older than previously described Miocene material and it represents the oldest known Neogene cordylids in Europe. The material of the dentary is very similar to that of Palaeocordylus bohemicus Roček, 1984, described from the younger locality of Dolnice near Cheb. The intramandibular septum of the dentary described herein, lying ventromedially from the alveolar canal, has a free posterolateral portion. It represents the first report of this structure in cordylids. The maxilla KEY WORDS Squamata, is only fragmentarily preserved. The size difference between the maxillary and Scincoidea, dentary elements, the number of teeth and, especially their morphology evokes Cordylidae, the possibility of two different taxa. Unfortunately, it is impossible to decide Palaeocordylus, Eggenburgian, on the basis of such limited material if both finds represent independent forms North-West Bohemia. or only extreme forms of the same taxon. GEODIVERSITAS • 2012 • 34 (4) © Publications Scientifiques du Muséum national d’Histoire naturelle, Paris. www.geodiversitas.com 837 Čerňanský A. RÉSUMÉ Les plus anciens « lézards » (Squamata, Cordylidae) du Néogène d’Europe. Remarques sur les immigrations de taxons africains au Miocène inférieur. Les plus anciens lézards cordylidés du Néogène européen proviennent du Miocène inférieur de Merkur-Nord (République Tchèque). L’histoire des cordylidés fossiles est très mal connue et ce groupe est rare dans le Miocène inférieur d’Europe. Les fossiles décrits ici proviennent des marnes calcaires grises de la base de la « Main Brown Coal Seam ». Ces marnes sont interprétées comme des cendres volcaniques remaniées. Les sédiments sont datés du Miocène inférieur (zone MN 3a). Ainsi, la faune de ce gisement est plus ancienne que celles décrites précédemment du Miocène. Le dentaire de ce cordylidé est similaire à celui de Palaeocordylus bohemicus Roček, 1984, provenant du gisement plus récent de Dolnice, situé près de Cheb. La portion postérolatérale du septum intramandi- MOTS CLÉS bulaire du dentaire, situé ventralement au canal alvéolaire, est libre. Le maxillaire Squamata, Scincoidea, est fragmentaire. La grande différence de taille entre maxillaire et dentaire, le Cordylidae, nombre de dents et, surtout, leur morphologie suggèrent la possibilité de deux Palaeocordylus, taxons différents. Malheureusement, sur la base du peu de matériel disponible, Eggenburgien, Bohême nord- il est impossible d’établir si ces fossiles représentent des formes distinctes ou occidentale. seulement des variations extrêmes du même taxon. INTRODUCTION or Ethiopia, respectively (Broadley & Branch 2002). Cordylidae contains 80 nominal taxa. This This paper deals with the first occurrence of the family has traditionally been divided into four family Cordylidae at the Lower Miocene (MN 3a) genera: Cordylus Gronovius, 1763, Pseudocordylus locality of Merkur-North. Girdled lizards (Cor- Smith, 1838, Chamaesaura Schneider, 1801 and dylidae) are sub-Saharan Africa’s only endemic Platysaurus Smith, 1844 (Loveridge 1944; Lang squamate family. This family belongs to a clade 1991). A recent molecular study has produced a of scincomorph lizards, the Cordyliformes. In new view of the cordylids. Stanley et al. (2010) addition to this family, this clade includes the used three nuclear and three mitochondrial genes Gerrhosauridae. The monophyly of cordyliformes from 111 specimens, representing 51 of the 80 based on morphological data suggested by Lang known taxa and recovered a comb-like tree with (1991) was confirmed by the cytogenetic study 10, well-supported, monophyletic lineages. Their and analysis of mitochondrial DNA by Odierna taxonomic reassessment divides the family into et al. (2002). Relationships to other lizards re- 10 genera that correspond to the well-supported main problematic, but they appear to be close lineages. They recovered evidence that supports two to the Scincidae (Estes et al. 1988; Lee 1998, subfamilies within the Cordylidae, the egg-laying 2002; Vicario et al. 2003; Whiting et al. 2003; Platysaurinae and the live-bearing Cordylinae. Townsend et al. 2004). The fossil record of cordylids is very poor. The Today, Cordylidae is the only lizard family oldest taxon is represented by a recently discovered restricted to Africa, and most species of the fam- fossilized lizard, Konkasaurus mahalana Krause, Ev- ily occur in South Africa and adjacent countries ans & Gao, 2003. This specimen has been assigned (e.g., Estes 1983; Branch 1984); only Cordylus to the Cordylidae, and it suggests that the family’s angolensis Bocage, 1895 and C. rivae (Boulenger, range once extended to Madagascar (Krause et al. 1896) can be found further north around Angola 2003). Assuming that the dating and taxonomy 838 GEODIVERSITAS • 2012 • 34 (4) The oldest known European Neogene girdled lizard fauna of K. mahalana is correct, the Cordylidae family is at least 68 million years old. The taxon Pseudolac­ Merkur-North erta mucronata (Filhol, 1877) from the Eocene of France (Augé 2003), formed the basis of the re- Prague cord of the Cordylidae in Europe. However, Augé (2005) removed the species P. lamandini (Filhol, CZECH REPUBLIC 1877) from Pseudolacerta De Stefano, 1903 and transferred it to the iguanid Geiseltaliellus Kuhn, 1944. He retained the species P. mucronata in Pseudolacerta (to which he added his new species 75 km P. quercyini Augé, 2005) and he assigned Pseu­ dolacerta to the Iguanidae. The last occurrence of FIG. 1. — Location of the Merkur-North locality, Czech Republic. cordylids in Europe is regarded as early Oligocene (Augé 2005). However, Böhme & Lang (1991) demonstrated that “Lacerta” rottensis von Meyer, All material was collected by screen-washing. The 1856 (latest Oligocene, MP30; Germany) should specimen is housed in the Geological collection of be referred to Cordyliformes, i.e. to the Cordylidae the Bilína opencast mine situated near Chomutov (Estes et al. 1988). in the western part of the Czech Republic. The Cordylids were rare during the Neogene of anatomical terminology of the individual skeletal Europe. Previously, Neogene cordylids from Eu- elements used here is mostly that of Féjervary-Langh rope were only known from Dolnice near Cheb (1923) and Roček (1984). The nomenclature of the (Palaeocordylus bohemicus Roček, 1984; Czech tooth crown follows Richter (1994) and Kosma Republic, MN 4), Obergänserndorf (Cordylidae (2004). The specimens were photographed using indet. Böhme 2002; Austria, MN5), Petersbuch 2 a scanning electron microscope (SEM). (Bavaricordylus ornatus Kosma, 2004; Germany, The Merkur-North locality (Lower Miocene, MN4a – however, this name was established in an MN 3a) is located in NW Bohemia and it was unpublished work – thesis, therefore it should be a discovered in an opencast (lignite) mine near Cho- nomen dubium; Rage pers. comm.), Puttenhausen mutov. The specimens are preserved in grey calcare- (Cordylidae indet. Abdul-Aziz et al. 2008; Ger- ous marls at the base of the so-called “Main Brown many, MN5 and the material described by Böhme Coal Seam”. The marls are interpreted as reworked (2010) as a new taxon of the genus Bavaricordylus – volcanic ash. This locality has already yielded a rich B. mollasicus; type locality is Puttenhausen B) and material of various groups of fish, amphibians and Sandelzhausen (MN 5; Böhme 2010). reptiles, e.g., frogs (Kvaček et al. 2004), albanertpe- The new material of cordylid lizards as described tontids (Čerňanský 2010a), lacertids (Čerňanský & below from the Lower Miocene locality of Merkur- Joniak 2009), chameleonids (Fejfar & Schleich North (MN 3a) is much older than previously 1994; Čerňanský 2010b), choristoderans (Evans & described European Neogene specimens and new Klembara 2005), amphisbaenids (Čerňanský & information about the oldest known Neogene Venczel 2011), gekkonids (Čerňanský & Bauer cordylids in Europe is given. 2010), anguimorph lizards (Klembara 2008) and snakes (Ivanov 2002). The sediments are also richly fossiliferous in remains of limnic and terrestrial MATERIAL, LOCALITY molluscs, plants and mammals (Fejfar et al. 1997a, AND GEOLOGICAL SETTING b, 1998; Kvaček et al. 2004). This study presents the first record of a cordylid lizard ABBREVIATIONS OF REPOSITORIES from the Lower Miocene (Eggenburgian, MN3a) Ah-number SGDB Geological collection