Translator Disclaimer
1 December 2010 Euleptes gallica Müller (Squamata: Gekkota: Sphaerodactylidae) from the Lower Miocene of North-West Bohemia, Czech Republic
Author Affiliations +

Gekkotan lizards are widely distributed in tropical regions and extend into more northern regions in Asia and Europe. Unfortunately, gekkotans lack significant fossil record and many aspects of their evolutionary biology are still poorly understood. Our study provides the first information about occurrence of this group in the Czech Republic. The fossil material comes from the Lower Miocene sediments (MN 3) of the locality Merkur-North. All the gekkonid bones can be assigned to the family Sphaerodactylidae and represent a single taxon, Euleptes gallica. This taxon was previously known only from the French locality Montaigu (zone MN 2), and thus our study extends its stratigraphic and geographic range.


The fossil record of gekkotan lizards is, in general, poor (Estes 1983, Augé 2005). Other than a few remains from Paleogene and Neogene deposits, most Cenozoic finds are no older than Pleistocene (Müller & Mödden 2001). Moreover, most of the fossil material found is very fragmentary, often rendering determination below the familial level impossible (Augé & Rage 2000, Augé 2005). A notable exception is the gecko found in Eocene Baltic amber (Bauer et al. 2005). The record of gekkotans from the Lower and Middle Miocene of Europe comprises just several remains of dentaries and maxillae and other isolated, often fragmentary, elements from Slovakia (Estes 1969), France (Hoffstetter 1946, Müller 2001, Augé et al. 2002), Germany (Schleich 1985, 1987, Müller & Mödden 2001), and Spain (Alferez Delgado & Brea Lopez 1981). Because of this limited material, the diversity and evolution of gekkotans in Europe remain largely unknown (Augé 2005). We here describe new material, referrable to the sphaerodactylid gecko Euleptes gallica from the lower Miocene locality of Merkur-North (zone MN 3) in the Czech Republic. This taxon was previously known only from the French locality Montaigu (zone MN 2).

Study Area

The locality is represented as an opencast mine near Chomutov. The specimens are preserved in grey calcareous marls at the base of the so-called, “Main Brown Coal Seam,” which are interpreted as reworked volcanic ash. This second Lower Miocene (MN 3) locality lies in the North West Bohemian rift and besides of gekkotan lizards yielded a rich material of various groups of amphibians and reptiles, e.g. frogs (Vejvalka 1997, Kvaček et al. 2004), chamaeleonids (Fejfar & Schleich 1994, Čerňanský 2010), lacertids (Čerňanský & Joniak 2009), an amphisbaenid lizard (Čerňanský & Venczel 2010), choristoderans (Evans & Klembara 2005), an anguiomorph lizard (Klembara 2008) and snakes (Ivanov 2002). The sediments are also richly fossiliferous in remains of limnic and terrestrial mollusks, plants, and mammals (Fejfar & Kvaček 1993, Fejfar et al. 1997a, b, 1998, Kvaček et al. 2004).

Material and Methods

This study is based on fossils housed in the Geological collection of the Bílina opencast mine, and consists of isolated elements collected by screen-washing or surface prospecting from the Lower Miocene (Eggenburgian), zone MN 3, of the Merkur-North locality, Czech Republic. The material was compared to the type material of Euleptes gallica and to other European fossil finds of the genus Euleptes. The material described by Schleich (1985, 1987) and Hofstetter (1946) can be excluded from a detailed comparison as they are not referable to the genus Euleptes. The first taxon, Palaeogekko risgoviensis Schleich, 1987 differs from the Czech gekkonid in lacking the triangular process at the anterior part of the maxilla. The jaw material of Gerandogekko described by Hofstetter (1946) is very poorly preserved. The majority of this material consists of vertebrae and one frontal. Unfortunately, these elements are unknown from the Czech locality. Likewise, material from the Middle Miocene of the French locality of Sansan (Augé & Rage 2000) is too fragmentary for an exact determination. Comparisons were also made with the sole extant member of the genus, E. europaea, based on a cleared and stained specimen (Texas Cooperative Wildlife Collection TCWC 78071), several radiographs (Museo Zoologico de la Specola, Universita di Firenze, MZUF 19119–19139) and published descriptions and illustrations (Wiedersheim 1875, Estes 1969, Rieppel & Schneider 1981). Osteological terminology follows that of Daza et al. (2009). The term supradental shelf for the maxilla (or subdental shelf for dentary) is widely used in the literature. Unfortunately, this name is ambiguously defined (see Estes et al. 1988, Conrad 2008, Rage & Augé 2010). Here, the term supradental shelf of maxilla is applied to a medial extension of the shelf that projects medially to the teeth and does not support them. All the gekkonid bones can be assigned to the single taxon, E. gallica, on the basis of a combination of features characteristic of this taxon and exhibited by the type material described from France by Müller (2001). Abbreviations of repositories: Ah-number SGDB, geological collection of the Bílina opencast mine, Czech Republic.


Euleptes gallica Müller, 2001

  • Locality and horizon: Merkur-North locality in Czech Republic, Lower Miocene (Eggenburgian), zone MN 3.

  • Material: Ah-875 SGDB, right dentary; Ah-931 SGDB, 932 SGDB, 933 SGDB, left maxillae.

  • Description: The dentary is a long, slender, elongate bone (Fig. 1), with a slight medial curvature at its anterior end. It is mediolaterally compressed and in lateral view heightens gradually posteriorly. The posteriormost portion of the element and the symphysial facet for the complementary dentary are missing. A high straight alveolar ridge supports a single row of at least 21 (probably more given the incomplete posterior margin of the element), typically uncuspid, pleurodont teeth (five teeth and 16 unoccupied tooth loci are present in Ah-875 SGDB). The alveolar shelf is proximately 50% higher than ventral portion of the dentary. The height of the shelf is identical along its whole length. A sulcus dentalis is present along the tooth row. In medial view, the closed Meckelian canal is exposed on the ventral side in the posterior third of the bone. The otherwise smooth labial surface of the dentary is pierced along its central portion by a longitudinal series of four mental foramina.

    The maxilla is subtriangular in shape (Fig. 2A). It is a rather elongate, relatively lightly-built bone. The highest portion is approximately 25% of the lenght of the entire maxilla. The nasal process (pars nasalis) is well-preserved in Ah-931 SGDB and is high and trapezoidal in shape. The anterior margin of the nasal process is thickened and slopes gently anteroventrally in a nearly straight line. Its terminus is free and well bounded from the dental portion (pars dentalis) of the maxilla and forms a triangular tip, typical for Euleptes spp. The ventral margin of the tip is oriented at 90° to the anterior margin of the maxilla. The posterior margin of the nasal process is higher and slopes more steeply than the anterior. The medial surface of the pars nasalis bears a posterodorsally directed curved ridge, the carina maxillaris (sensu Müller 1996), which contributes to the anterolateral wall of the nasal capsule. This feature terminates in a narrowing, posteroventrally-arching branch near the center of the pars nasalis, dorsal and parallel to a less prominent branch of the carina maxillaris. Twentyone tooth loci are present in Ah-931 SGDB and 20 in Ah-932 SGDB. The alveolar ridge is straight and high. The supradental shelf (lamina horizontalis) is very well developed and widened, particulary in its anterior portion. In the middle portion of the shelf there is a prominent horizontal expansion (the palatine process of maxilla), the medial edge of which is slightly recurved dorsaly. Its edge is thickened, narrowing as the shelf decreases in width posteriorly. In Ah-932 SGDB a well preserved infraorbital foramen lies above the medial shelf, approximately at its midpoint (Fig. 2B). The posterior portion of the maxillary is slender and straight, and tapers slightly distally. It bears an elongated jugal facet.

    The otherwise smooth labial surface of the maxilla is pierced along its lower margin by a series of ten supralabial foramina, whereas a shorter series of three foramina lie dorsal to these.

  • Dentition: A high number of pleurodont teeth is the classic feature of the gekkotan lizards. The sizes of inter-dental gaps are very small. The teeth are slender, smooth, conical and straight, with those in the central portion of the dentary highest. Small circular resorption pits are present on the lingual sides of the tooth bases. In general, tooth shape of maxilla is similar to that of the dentary.

  • Comparisons: The Czech material described here is referable to the genus Euleptes on the basis of the triangular tip of the anterior margin of the pars nasalis of the maxillary and the smooth dorsal margin of the pars nasalis, lacking a pronounced dorsal process (Bauer et al. 1997, Müller & Mödden 2001). This configuration of the nasal process contrasts sharply with the prominent, often pointed process typical of many gekkotans (Daza 2008), including the Middle Miocene Palaeogekko risgoviensis (Schleich 1987). The specifically diagnostic features of the maxilla are those recorded for E. gallica by Müller (2001) and the fragmentary dentary is, likewise, fully compatible with that of this species. We, therefore, refer the new material to E. gallica.

    Euleptes gallica differs from the extant Euleptes europaea in having a straight (versus weakly concave [in our comparative material, although illustrated by Estes (1969) as weakly convex]) anterior margin of the pars nasalis of the maxilla, terminating in a more prominent and more anteriorly-projecting triangular tip. Further, although our material is indicative of smaller specimens (estimated SVL 60–70 mm) than those (estimated SVL 85 mm) reported by Müller (2001), this is still much larger than the living species, which reaches a maximum of only 45 mm SVL (Delaugerre 1985; although Estes (1969) gives a SVL of „approximately 50 mm“).

    Our material of E. gallica differs from Phyllodactylus (=Euleptes) sp. from the middle Miocene of Slovakia described by Estes (1969) by a distinctly lower dorsal process of the maxilla and a greater number of supralabial foramina (13 in two rows versus seven).

    Our material may be distinguished from Euleptes sp. described from Germany (Müller & Mödden 2001) by its more gently sloping and straighter (versus steeply angled and weakly concave) anterior margin of the pars nasalis of the maxilla, and the presence of a large and prominent carina maxillaris on the medial surface of the nasal process of the maxilla (described as small in Euleptes sp. by Müller & Mödden (2001)). Unfortunately, the fragmentary dentary does not provide diagnostic information relevant at the species level. Likewise tooth size, shape, and number appear to be consistent across Euleptes spp. We estimate the total number of maxillary tooth loci at 28 in the Czech material of E. gallica. Estes reported 29 loci for Euleptes sp. from Slovakia, and Müller & Möden (2001) reported 26–28 in Euleptes sp. from Germany. Bauer et al. (1997) reported approximately 25 maxillary teeth in extant E. europaea, but we have confirmed that this may reach 30 loci in some specimens (e.g., TCWC 78071). Müller (2001) noted at least 31 loci in the larger type material of E. gallica. Given typical intraspecific variation in tooth counts among gekkotans and the increase in number of loci with body size within a single species (Bauer & Russell 1990) we consider this character as uninformative.

  • Fig. 1.

    Euleptes gallica: the right dentary Ah-875 SGDB, in medial view.


    Fig. 2.

    Euleptes gallica: A) the left maxilla Ah-931 SGDB, in internal view, B) posterior ending of the left maxilla Ah-932 SGDB, in internal view.



    The European gecko genus Euleptes, represented by a single extant species as well as fossil material, has been long synonymized within a nearly cosmopolitan Phyllodactylus. Bauer et al. (1997) dismantled Phyllodactylus, leaving it restricted to the New World and resurrected Euleptes. The first approaches to partition Phyllodactylus were considerably earlier (Dixon & Kroll 1974). Recent higher order reclassification of gekkotan lizards based on molecular phylogenetics has unambiguously placed Euleptes within the family Sphaerodactylidae (Gamble et al. 2008), within which it is the only leaftoed genus. The Merkur-North material represents the first proof of the former existence of the family Sphaerodactylidae and, indeed, of the Gekkota in the Czech Republic. The finding seems noteworthy from three points. First, the new find thus demonstrates the occurence of the species E. gallica not only in western Europe, but in central Europe as well, and expands its stratigraphic range to somewhat younger deposits. Second, since the distribution of gekkotan lizards is confined to subtropical and tropical regions (Estes 1983), this finding, together with fossil remains of chameleons from the same locality (Fejfar & Schleich 1994, Čerňanský 2010), further corroborates evidence for tropical or subtropical climatic conditions in central Europe during the early Miocene (Böhme 2003). Finally, it contributes to our understanding of the paleodiversity and evolution of the family Sphaerodactylidae.


    We are indebted to J. Klembara (Comenius University in Bratislava) for the opportunity to study material described in this paper. T. Hibbits (TCWC) kindly provided Euleptes europaea for comparison. For critically reading the manuscript and the text corrections we thank Dr. M. Augé (Muséum National D`Histoire Naturelle Paris) and Dr. L. Gvoždík (Institute of Vertebrate Biology, Academy of Sciences of the Czech Republic, v.v.i.). This project was supported by APVV Company, G. No. 0280-07 (AC) and the National Science Foundation of the United States (DEB 0844523).



    Alferez Delgado F. & Brea Lopez P. 1981: Estudio preliminar de los restos de peces, anfibios y reptiles del yacimiento mioceno de Corcoles (Guadalajara). Bol. R. Soc. Esp. Hist. Nat. Secc. Geol. 79: 5–20. Google Scholar


    Augé M. 2005: Évolution des Lézards du Paléogène en Europe. Mèm. Mus. Natn. Hist. Nat. 192: 1–399. Google Scholar


    Augé M. & Rage J.C. 2000: Les Squamates (Reptilia) du Miocène moyen de Sansan. In: Ginsburg L. (ed.), La faune miocène de Sansan et son environnement. Mèm. Mus. Natn. Hist. Nat. 183: 263–313. Google Scholar


    Augé M., Ginsburg L., de Lapparent de Broin F., Makinsky M., Mourer C., Pouit D. & Sen S. 2002: Les vertébrés du Miocène moyen de Contres (Loir-et-Cher, France). Revue de Paléobiologie 21 (2) : 819–852.  Google Scholar


    Bauer A.M. & Russell A.P. 1990: Dentitional diversity in Rhacodactylus (Reptilia: Gekkonidae). Mem. Qld. Mus. 29: 311–322. Google Scholar


    Bauer A.M., Böhme W. & Weitschat W. 2005: An early Eocene gecko from baltic amber and its implications for the evolution of geko adhesion. J. Zool., London 265: 327–332. Google Scholar


    Bauer A.M., Good D.A. & Branch W.R. 1997: The taxonomy of the Southern African leaf-toed geckos (Squamata: Gekkonidae) with a review of Old World “Phyllodactylus” and the description of five new genera. Proc. Calif. Acad. Sci. 49: 447–497. Google Scholar


    Böhme M. 2003: The Miocene climatic optimum: Evidence from ectothermic vertebrates of Central Europe. Palaeogeography, Palaeoclimatology, Palaeoecology 195: 389–401. Google Scholar


    Conrad J.L. 2008: Phylogeny and systematics of Squamata (reptilia) based on morphology. Bull. Amer. Mus. Nat. Hist. 310: 1–182. Google Scholar


    Čerňanský A. 2010: A revision of chamaeleonids from the Lower Miocene of the Czech Republic with description of a new species of Chamaeleo (Squamata, Chamaeleonidae). Geobios Google Scholar


    Čerňanský A. & Joniak P. 2009: New finds of lacertids (Sauria, Lacertidae) from the Neogene of Slovakia and Czech. Acta Geologica Slovaca 1: 57–64. Google Scholar


    Čerňanský A. & Venczel M. 2010: An amphisbaenid reptile (Squamata, Amphisbaenidae) from the Lower Miocene of Northwest Bohemia (MN 3, Czech Republic). Neues Jahrbuch für Geologie und Paläeontologie - Abhandlungen (in press).  Google Scholar


    Daza J.D. 2008: Cladistic analysis of the Gekkota (Reptilia) by means of craniological data. Ph.D. dissertation, University of Puerto Rico, Rio PiedrasGoogle Scholar


    Daza J.D., Abdala V., Thomas R. & Bauer A.M. 2009: Skull anatomy of the miniaturized gecko Sphaerodactylus roosevelti (Squamata: Gekkota). J. Morphol. 269: 1340–1364. Google Scholar


    Delaugerre M. 1985: La variation géographique chez Phyllodactylus europaeus Gené (Reptilia, Sauria, Gekkonidae) etude de la population de l'Îlot Sperduto Grande (sud de la Corse, Reserve Naturelle des Îles Lavezzi). Bull. Mensuel de la Société Linnéenne de Lyon 54: 262–269. Google Scholar


    Dixon J.R. & Kroll J.C. 1974: Resurrection of the generic name Paroedura for the phyllodactyline geckos of Madagascar, and description of a new species. Copeia 1: 24–30. Google Scholar


    Estes R. 1969: Die Fauna der miozänen Spaltenfühlung von Neudorf an der March (CSSR) (Reptilia, Lacertilia). Österr. Akad. Wiss., Math.-Naturwiss. Kl. Abt. I, Sitz. Ber. 178: 77–82. Google Scholar


    Estes R. 1983: Encyclopedia of Paleoherpetology, Part 10 A-Sauria terrestria, Amphisbaenia. Gustav Fischer VerlagGoogle Scholar


    Estes R., de Queiroz K. & Gauthier J. 1988: Phylogenetic relationships within Squamata. In: Estes R. & Pregill G. (eds.), Phylogenetic relationships of the lizard Families. Stanford University Press : 119–281. Google Scholar


    Evans S. & Klembara J. 2005: A choristoderan reptile (Reptilia, Diapsida), from the Lower Miocene of Northwest Bohemia (Czech Republic). J. Vertebrate Paleontol. 25: 171–184. Google Scholar


    Fejfar O. & Kvaček Z. 1993: Excursion Nr. 3, Tertiary basins in Northwest Bohemia. Charles University, Czech Geological Society.  Google Scholar


    Fejfar O. & Schleich H.H. 1994: Ein Chamäleonfund aus dem unteren Orleanium des Braunkohlen-Tagebaus Merkur-Nord (Nordböhmen). Courier Forschungsinstitut Senckenberg 173: 167–173. Google Scholar


    Fejfar O., Engesser B. & Tomida Y. 1997a: New eomyid genus and species of Apeomys Falbusch (Eomyidae, Rodentia, Mammalia). In: Aguilar J.P., Legendre S. & Michaux J. (eds.), Actes du Congrès BiochroM '97. Mémoires et Travaux de l'Ecole Pratique des Hautes Études. Institut de Montpellier 21: 705–706. Google Scholar


    Fejfar O., Heizmann E.P.J. & Major P. 1997b: Metaschizotherium cf. wetzleri (Kowalewsky) from the early Miocene of the Czech Republic and South Germany. In: Aguilar J.P., Legendre S. & Michaux J. (eds.), Actes du Congrès BiochroM '97. Mémoires et Travaux de l'Ecole Pratique des Hautes Études. Institut de Montpellier 21: 707–709. Google Scholar


    Fejfar O., Rummel M. & Tomida Y. 1998: New eomyid genus and species from the early Miocene (MN zones 3–4) of Europe and Japan related to Apeomys (Eomyidae, Rodentia, Mammalia). In: Tomida I., Flynn L.J. & Jacobs L.L. (eds.), Advances in vertebrate paleontology and geochronology. National Science Museum Monographs 14: 123–143. Google Scholar


    Gamble T., Bauer A.M., Greenbaum E. & Jackman T.R. 2008: Evidence for Gondwanan vicariance in an ancient clade of gecko lizards. J. Biogeography 35: 88–104. Google Scholar


    Hoffstetter R. 1946: Sur les gekkonidae fossiles. Bull. Mus. Hist. Nat. Paris , 18: 195–203. Google Scholar


    Ivanov M. 2002: The oldest known Miocene fauna from Central Europe: Merkur-North locality, Czech Republic. Acta Paleontologica Polonica 47: 513–534. Google Scholar


    Klembara J. 2008: A new anguiomorph lizard from the Lower Miocene of North-West Bohemia, Czech Republic. Paleontology 51: 81–94. Google Scholar


    Kvaček Z., Böhme M., Dvořák Z., Konzalová M., Mach K., Prokop J. & Rajchl M. 2004: Early Miocene freshwater and swamp ecosystems of the Most basin (northern Bohemia) with particular reference to the Bílina mine section. J. Czech Geol. Soc. 49: 1–40. Google Scholar


    Müller J. 1996: Eine neue Art der echten Eidechsen (Reptilia: Lacertilia: Lacertidae) aus dem Unteren Miozän von Poncenat, Frankreich. Mainzer Geowiss. Mitt. 25: 79–88. Google Scholar


    Müller J. 2001: A new fossil species of Euleptes from early Miocene of Montaigu, France (Reptilia, Gekkonidae). Amphibia-Reptilia 22: 342–347. Google Scholar


    Müller J. & Mödden C. 2001: A fossil leaf-toed gecko from the Oppenheim/Nierstein Quarry (Lower Miocene, Germany). J. Herpetol. 35: 532–537. Google Scholar


    Rage J.C. & Augé M. 2010: Squamate reptiles from the middle Eocene of Lissieu (France). A landmark in the middle Eocene of Europe. Geobios 43: 243–268. Google Scholar


    Rieppel O. & Schneider B. 1981: Phyllodactylus europaeus Gené 1838 - Europäischer Blattfingergecko. In: Böhme W. (ed.), Handbuch der Reptilien und Amphibien Europas, Band 1, Echsen I. Akademische Verlagsgesellschaft , Wiesbaden : 108–118. Google Scholar


    Schleich H.H. 1985: Zur Verbreitung tertiärer und quartärer Reptilien und Amphibien. Müncher Geowiss. Abh. 4: 67–93. Google Scholar


    Schleich H.H. 1987: Neue Reptilienfunde aus dem Tertiär Deutschlands 7. Erstnachweis von Geckos aus dem Mittelmiozänen Süddeutchlands: Palaeogekko risgoviensis nov. gen., nov. spec. (Reptilia, Sauria, Gekkonidae). Mitt. Bayer. Staatsslg. Paläont. hist. Geol. 27: 67–93. Google Scholar


    Vejvalka J. 1997: Amphibians (Amphibia: Caudata, Salienta) and Reptiles (Reptilia: Lacertilia, Choristodera) of the Miocene locality Merkur-North (Czech Republic). Thesis, Faculty of Natural Sciences, Charles University in Prague.  Google Scholar


    Wiedersheim R. 1875: Zur Anatomie und Physiologie des Phyllodactylus europaeus, mit besonderer Berücksichtigung des Aquaeductus vestibuli der Ascalaboten im Allgemeinen. Zugleich als zweiter Beitrag zur Inselfauna des Mittelmeeres. Morph. Jb. 1: 495–534. Google Scholar
    Andrej Čerňanský and Aaron M. Bauer "Euleptes gallica Müller (Squamata: Gekkota: Sphaerodactylidae) from the Lower Miocene of North-West Bohemia, Czech Republic," Folia Zoologica 59(4), 323-328, (1 December 2010).
    Received: 20 January 2010; Accepted: 1 June 2010; Published: 1 December 2010
    6 PAGES

    Back to Top