Fossil lizards and worm lizards (Reptilia, Squamata) from the Neogene and Quaternary of Europe: an overview

Abstract

Lizards were and still are an important component of the European herpetofauna. The modern European lizard fauna started to set up in the Miocene and a rich fossil record is known from Neogene and Quaternary sites. At least 12 lizard and worm lizard families are represented in the European fossil record of the last 23 Ma. The record comprises more than 3000 occurrences from more than 800 localities, mainly of Miocene and Pleistocene age. By the beginning of the Neogene, a marked faunistic change is detectable compared to the lizard fossil record of Palaeogene Europe. This change is reflected by other squamates as well and might be related to an environmental deterioration occurring roughly at the Oligocene/Miocene boundary. Nevertheless, the diversity was still rather high in the Neogene and started to decrease with the onset of the Quaternary glacial cycles. This led to the current impoverished lizard fauna, with the southward range shrinking of the most thermophilic taxa (e.g., agamids, amphisbaenians) and the local disappearance of other groups (e.g., varanids). Our overview of the known fossil record of European Neogene and Quaternary lizards and worm lizards highlighted a substantial number of either unpublished or poorly known occurrences often referred to wastebasket taxa. A proper study of these and other remains, as well as a better sampling of poorly explored time ranges (e.g., Pliocene, Holocene), is needed and would be of utmost importance to better understand the evolutionary history of these reptiles in Europe.

This is a preview of subscription content, log in to check access.

Fig. 1
Fig. 2
Fig. 3
Fig. 4
Fig. 5
Fig. 6
Fig. 7
Fig. 8
Fig. 9
Fig. 10
Fig. 11
Fig. 12
Fig. 13
Fig. 14

References

  1. Abbazzi, L., Angelone, C., Arca, M., Barisone, G., Bedetti, C., Delfino, M., et al. (2004). Plio-Pleistocene fossil vertebrates of Monte Tuttavista (Orosei, Eastern Sardinia, Italy), an overview. Rivista Italiana di Paleontologia e Stratigrafia,110, 681–706.

    Google Scholar 

  2. Abdul Aziz, H., Böhme, M., Rocholl, A., Zwing, A., Prieto, L., Wijbrans, J. R., et al. (2008). Integrated stratigraphy and 40Ar/39Ar chronology of the early to middle Miocene Upper Freshwater Molasse in eastern Bavaria (Germany). International Journal of Earth Sciences,97, 115–134.

    Google Scholar 

  3. Adams, A. L. (1866). On fossil chelonians from the ossiferous caves and fissures of Malta. Quarterly Journal of the Geological Society,22, 594–595.

    Google Scholar 

  4. Agustí, J., Blain, H.-A., Cuenca-Bescòs, G., & Bailon, S. (2009). Climate forcing of first hominid dispersal in Western Europe. Journal of Human Evolution,57, 815–821.

    Google Scholar 

  5. Albert, E. M., & Fernández, A. (2009). Evidence of cryptic speciation in a fossorial reptile: description of a new species of Blanus (Squamata: Amphisbaenia: Blanidae) from the Iberian Peninsula. Zootaxa,2234, 56–68.

    Google Scholar 

  6. Alexejew, A. (1912). Description de la faune méotique des vertébrés des environ du village Petroviérovak (District Tiraspol). I. Anguidae. Zapiski matematicheskogo otdeleniya Novorossiiskogo obshchestva estestvoispytatelei,39, 13–40.

    Google Scholar 

  7. Arnold, N., & Ovenden, D. (2002). A field guide to the reptiles and amphibians of Britain and Europe. London: Harper-Collins Publisher.

    Google Scholar 

  8. Augé, M. (1993). Une nouvelle espèce de Lacertidé (Reptilia, Lacertilia) des Faluns Miocènes de l’Anjou-Touraine. Bulletin de la Société de Sciences naturelles de l’Ouest de la France,15, 69–74.

    Google Scholar 

  9. Augé, M. (2003). La faune de Lacertilia (Reptilia, Squamata) de l’Éocène inférieur de Prémontré (Bassin de Paris, France). Geodiversitas,25, 539–574.

    Google Scholar 

  10. Augé, M. (2005). Évolution des lézards du Paléogène en Europe. Mémoires du Muséum national d’Histoire naturelle,192, 1–369.

    Google Scholar 

  11. Augé, M. (2012). Amphisbaenians from the European Eocene: a biogeographical review. Palaeobiodiversity and Palaeoenvironments,92, 425–443.

    Google Scholar 

  12. Augé, M., Bailon, S., & Malfay, J. P. (2003). Un nouveau genre de lacertidae (Reptilia, Lacertilia) dans les faluns miocènes de l’Anjou-Touraine (Maine-et-Loire, France). Geodiversitas,25, 289–295.

    Google Scholar 

  13. Augé, M., & Pouit, D. (2012). Presence of iguanid lizards in the European Oligocene. Lazarus taxa and fossil abundance. Bulletin de la Société Géologique de France,183, 653–660.

    Google Scholar 

  14. Augé, M., & Rage, J.-C. (2000). Les squamates (Reptilia) du Miocène moyen de Sansan. In L. Ginsburg (Ed.), La faune miocène de Sansan et son environnement. Mémoires du Muséum national d’Histoire naturelle, 183, 263–313.

  15. Augé, M., & Smith, R. (2009). An assemblage of early Oligocene lizards (Squamata) from the locality of Boutersem (Belgium), with comments on the Eocene-Oligocene transition. Zoological Journal of the Linnean Society,155, 148–170.

    Google Scholar 

  16. Augé, M., & Sullivan, R. M. (2006). A new genus, Paraplacosauriops (Squamata, Anguidae, Glyptosaurinae), from the Eocene of France. Journal of Vertebrate Paleontology,26, 133–137.

    Google Scholar 

  17. Bailon, S. (1991). Amphibiens et reptiles du Pliocène et du Quaternaire de France et d’Espagne: mise en place et evolution des faunes. Paris: Université Paris VII.

    Google Scholar 

  18. Bailon, S. (2004). Fossil record of Lacertidae in Mediterranean islands: the state of the art. In V. Pérez-Mellado, N. Riera, & A. Perera (Eds.), The biology of lacertid lizards. Evolutionary and ecological perspectives. Institut Menorquí d’Estudis, Recerca, 8, 37–62.

  19. Bailon, S., & Augé, M. (2012). Un nouveau genre, Ragesaurus (Squamata, Anguidae, Anguinae), du Pléistocène inférieur des îles Medas (Catalogne, Espagne). Bulletin de la Societe Geologique de France,183, 683–688.

    Google Scholar 

  20. Bailon, S., & Blain, H.-A. (2007). Faunes de reptiles et changements climatiques en Europe occidentale autour de la limite Plio-Pléistocène. Quaternaire,18, 55–63.

    Google Scholar 

  21. Bailon, S., Boistel, R., Bover, P., & Alcover, J. A. (2014). Maioricalacerta rafelinensis, gen. et sp. nov., (Squamata, Lacertidae), from the early Pliocene of Mallorca (Balearic Islands, Western Mediterranean Sea). Journal of Vertebrate Paleontology,34, 318–326.

    Google Scholar 

  22. Barahona, F., & Barbadillo, L. J. (1997). Identification of some Iberian lacertids using skull characters. Revista Española de Herpetología,11, 47–62.

    Google Scholar 

  23. Barbadillo, L. J., García-París, M., & Sanchiz, B. (1997). Orígenes y relaciones evolutivas de la herpetofauna ibérica. In J. M. Pleguezuelos (Ed.), Distribución y biogeografía de los anfibios y reptiles en España y Portugal. Monografías de Herpetología, 3, 47–100.

  24. Bell, C. J., Gauthier, J. A., & Bever, G. S. (2010). Covert biases, circularity, and apomorphies: a critical look at the North American Quaternary Herpetofaunal Stability Hypothesis. Quaternary International,217, 30–36.

    Google Scholar 

  25. Blain, H.-A. (2005). Contribution de la paleoherpetofaune (Amphibia & Squamata) a la connaissance de l’evolution du climat et du paysage du Pliocene superieur au Pleistocene Moyen d’Espagne. Paris: Muséum national d’Histoire naturelle.

    Google Scholar 

  26. Blain, H.-A. (2009). Contribution de la paléoherpétofaune (Amphibia & Squamata) à la connaissance de l’évolution du climat et du paysage du Pliocène supérieur au Pléistocène moyen d’Espagne. Treballs del Museu de Geologia de Barcelona,16, 39–170.

    Google Scholar 

  27. Blain, H.-A. (2015). Anfibios y escamosos de Cueva Victoria. - Amphibians and squamate reptiles from Cueva Victoria. Mastia, 11-12-13, 175–197.

  28. Blain, H.-A., Agustí, J., Lordkipanidze, D., Rook, L., & Delfino, M. (2014a). Paleoclimatic and paleoenvironmental context of the Early Pleistocene hominins from Dmanisi (Georgia, Lesser Caucasus) inferred from the herpetofaunal assemblage. Quaternary Science Reviews,105, 136–150.

    Google Scholar 

  29. Blain, H.-A., & Bailon, S. (2010). Anfibios y escamosos del Pleistoceno inferior de Barranco León y de Fuente Nueva 3 (Orce, Andalucía, España). In I. Toro, B. Martínez-Navarro, & J. Agustí (Eds.), Ocupaciones humanas en el Pleistoceno inferior y medio de la Cuenca de Guadix-Baza (pp. 165–183). Sevilla: Consejería de Cultura.

    Google Scholar 

  30. Blain, H.-A., Bailon, S., & Agustí, J. (2007). Anurans and squamate reptiles from the latest early Pleistocene of Almenara-Casablanca-3 (Castellón, East of Spain). Systematic, climatic and environmental considerations. Geodiversitas,29, 269–295.

    Google Scholar 

  31. Blain, H.-A., Bailon, S., & Agustí, J. (2008). Amphibians and squamate reptiles from the latest Early Pleistocene of Cueva Victoria (Murcia, southeastern Spain, SW Mediterranean): paleobiogeographic and paleoclimatic implications. Geologica Acta,6, 345–361.

    Google Scholar 

  32. Blain, H.-A., Bailon, S., & Agustí, J. (2016). The geographical and chronological pattern of herpetofaunal Pleistocene extinctions on the Iberian Peninsula. Comptes Rendus Palevol,15, 761–764.

    Google Scholar 

  33. Blain, H.-A., Bailon, S., Cuenca-Bescós, G., Arsuaga, J. L., Bermúdez De Castro, J. M., & Carbonell, E. (2009). Long-term climate record inferred from early-middle Pleistocene amphibian and squamate reptile assemblages at the Gran Dolina Cave. Atapuerca. Spain. Journal of Human Evolution,56, 55–65.

    Google Scholar 

  34. Blain, H.-A., Gleed-Owen, C. P., López-García, J. M., Carrión, J. S., Jennings, R., Finlayson, G., et al. (2013). Climatic conditions for the last Neanderthals: herpetofaunal record of Gorham’s cave. Gibraltar. Journal of Human Evolution,64, 289–299.

    Google Scholar 

  35. Blain, H.-A., Lozano-Fernández, I., Ollé, A., Rodríguez, J., Santonja, M., & Pérez-González, A. (2015). The continental record of Marine Isotope Stage 11 (Middle Pleistocene) on the Iberian Peninsula characterized by herpetofaunal assemblages. Journal of Quaternary Science,30, 667–678.

    Google Scholar 

  36. Blain, H.-A., Panera, J., Uribelarrea, D., Rubio-Jara, S., & Pérez-González, A. (2012). Characterization of a rapid climate shift at the MIS 8/7 transition in central Spain (Valdocarros II, Autonomous Region of Madrid) by means of the herpetological assemblages. Quaternary Science Reviews,47, 73–81.

    Google Scholar 

  37. Blain, H.-A., Santonja, M., Pérez-González, A., Panera, J., & Rubio-Jara, S. (2014b). Climate and environments during Marine Isotope Stage 11 in the central Iberian Peninsula: the herpetofaunal assemblage from the Acheulean site of Áridos-1, Madrid. Quaternary Science Reviews,94, 7–21.

    Google Scholar 

  38. Bloos, G., Böttcher, R., Heinrich, W.-D., & Münzing, K. (1991). Ein Vorkommen von Kleinvertebraten in jung-Pleistozänen Deckschichten (Wende Eem/Würm) bei Steinheim an der Murr. Suttgarter Beiträge zur Naturkunde. Serie B (Geologie und Paläontologie),179, 1–72.

    Google Scholar 

  39. Boettger, O. (1875). Über die Gliederung der Cyrenenmergel-Gruppe im Mainzer Becken. Bericht über die Senckenbergische Naturforschende Geselschaft,1873–1874, 50–102.

    Google Scholar 

  40. Boettger, O. (1876/1877). Die Fauna der Corbicula-Schichten im Mainzer Becken. Palaeontographica, 24, 185–219.

  41. Böhme, M. (1999a). Die Miozäne Fossil-Lagerstätte Sandelzhausen. 16. Fisch-und Herpetofauna - Erste Ergebnisse. Neues Jahrbuch für Geologie und Paläontologie, Abhandlungen,214, 487–496.

    Google Scholar 

  42. Böhme, M. (1999b). Doppelschleichen (Sauria, Amphisbaenidae) aus dem Untermiozän von Stubersheim 3 (Süddeutschland). Mitteilungen der Bayerischen Staatssammlung für Paläontologie und Historische Geologie,39, 85–90.

    Google Scholar 

  43. Böhme, M. (2002). Lower vertebrates (Teleostei, Amphibia, Sauria) from the Karpatian of the Korneuburg Basin - palaeoecological, environmental and palaeoclimatical implications. Beiträge zur Paläontologie,27, 339–353.

    Google Scholar 

  44. Böhme, M. (2003). The Miocene Climatic Optimum: evidence from ectothermic vertebrates of Central Europe. Palaeogeography, Palaeoclimatology, Palaeoecology,195, 389–401.

    Google Scholar 

  45. Böhme, M. (2010). Ectothermic vertebrates (Actinopterygii, Allocaudata, Urodela, Anura, Crocodylia, Squamata) from the Miocene of Sandelzhausen (Germany, Bavaria) and their implications for environment reconstruction and palaeoclimate. Paläontologische Zeitschrift,84, 3–41.

    Google Scholar 

  46. Böhme, M., & Ilg, A. (2003). fosFARbase www.wahre-staerke.com/Accessed in 2017.

  47. Böhme, M., & Vasilyan, D. (2014). Ectotermic vertebrates from the late middle Miocene of Gratkorn (Austria, Styria). Palaeobiodiversity and Palaeoenvironments,94, 21–40.

    Google Scholar 

  48. Böhme, W., & Zammit-Maempel, G. (1982). Lacerta siculimelitensis sp. n. (Sauria: Lacertidae), a giant lizard from the Late Pleistocene of Malta. Amphibia-Reptilia,3, 257–268.

    Google Scholar 

  49. Bolet, A. (2017). First early Eocene lizards from Spain and a study of the compositional changes between late Mesozoic and early Cenozoic Iberian lizard assemblages. Palaeontologia Electronica, 20.2.20A, 1–22.

  50. Bolet, A., Daza, J. D., Augé, M., & Bauer, A. M. (2015). New genus and species names for the Eocene lizard Cadurcogekko rugosus Augé, 2005. Zootaxa,3985, 265–274.

    Google Scholar 

  51. Bolet, A., Delfino, M., Fortuny, J., Almécija, S., Robles, J. M., & Alba, D. M. (2014). An amphisbaenian skull from the European Miocene and the evolution of Mediterranean worm Lizards. PLoS ONE,9, e98082.

    Google Scholar 

  52. Bolet, A., & Evans, S. E. (2013a). Fossil history of chamaeleons. In K. A. Tolley & A. Herrel (Eds.), The biology of chamaeleons (pp. 175–192). Berkeley: University of California Press.

    Google Scholar 

  53. Bolet, A., & Evans, S. E. (2013b). Lizards and amphisbaenians (Reptilia, Squamata) from the late Eocene of Sossís (Catalonia, Spain). Palaeontologia Electronica, 16.1.8A, 1–23.

  54. Bolkay, S. J. (1913). Additions to the fossil herpetology of Hungary from the Pannonian and Praeglacial periode. Jahrbuche der Ungarischen geologischen Reichsanstalt,21, 217–230.

    Google Scholar 

  55. Bonfiglio, L., & Insacco, G. (1992). Palaeoenvironmental, paleontologic and stratigraphic significance of vertebrate remains in Pleistocene limnic and alluvial deposits from Southeastern Sicily. Palaeogeography, Palaeoclimatology, Palaeoecology,95, 195–208.

    Google Scholar 

  56. Bonfiglio, L., Marra, A. C., Masini, F., Pavia, M., & Petruso, D. (2002). Pleistocene faunas of Sicily: a review. In W. H. Waldren, & J. A. Ensenyat (Eds.), World islands in prehistory: international insular investigations. British Archaeological Reports, International Series, 1095, 428–436.

  57. Brunner, G. (1954). Das Fuchsloch bei Siegmannsbrunn (Oberfr.) (Eine mediterrane Riss-Wiirm-Fauna). Neues Jahrbuch für Geologie und Palaontologie,100, 83–118.

    Google Scholar 

  58. Caldwell, M. W., Nydam, R. L., Palci, A., & Apesteguía, S. (2015). The oldest known snakes from the Middle Jurassic-Lower Cretaceous provide insights on snake evolution. Nature Communications,6, 5996.

    Google Scholar 

  59. Caloi, L., Kotsakis, T., & Palombo, M. R. (1986). La fauna a vertebrati terrestri del Pleistocene delle isole del Mediterraneo. Geologica Romana,25, 235–256.

    Google Scholar 

  60. Carranza, S., & Arnold, E. N. (2006). Systematics, biogeography, and evolution of Hemidactylus geckos (Reptilia: Gekkonidae) elucidated using mitochondrial DNA sequences. Molecular Phylogenetics and Evolution,38, 531–545.

    Google Scholar 

  61. Ceríaco, L. M. P., & Bauer, A. M. (2018). An integrative approach to the nomenclature and taxonomic status of the genus Blanus Wagler, 1830 (Squamata: Blanidae) from the Iberian Peninsula. Journal of Natural History,52, 849–880.

    Google Scholar 

  62. Čerňanský, A. (2010a). A revision of chamaeleonids from the lower Miocene of the Czech Republic with description of a new species of Chamaeleo (Squamata, Chamaeleonidae). Geobios,43, 605–613.

    Google Scholar 

  63. Čerňanský, A. (2010b). Earliest world record of green lizards (Lacertilia, Lacertidae) from the Lower Miocene of Central Europe. Biologia,65, 737–741.

    Google Scholar 

  64. Čerňanský, A. (2011). A revision of the chameleon species Chamaeleo pfeili Schleich (Squamata; Chamaeleonidae) with description of a new material of chamaeleonids from the Miocene deposits of southern Germany. Bulletin of Geosciences,86, 275–282.

    Google Scholar 

  65. Čerňanský, A. (2012). The oldest known European Neogene girdled lizard fauna (Squamata, Cordylidae), with comments on early Miocene immigration of African taxa. Geodiversitas,34, 837–848.

    Google Scholar 

  66. Čerňanský, A. (2016). Another piece of the puzzle: the first report on the Early Miocene lizard fauna from Austria (Ottnangian, MN 4; Oberdorf locality). Paläontologische Zeitschrift,90, 723–746.

    Google Scholar 

  67. Čerňanský, A., & Bauer, A. M. (2010). Euleptes gallica Müller (Squamata: Gekkota: Sphaerodactylidae) from the Lower Miocene of North-West Bohemia, Czech Republic. Folia Zoologica,59, 323–328.

    Google Scholar 

  68. Čerňanský, A., Bolet, A., Müller, J., Rage, J.-C., Augé, M., & Herrel, A. (2017a). A new exceptionally preserved specimen of Dracaenosaurus (Squamata, Lacertidae) from the Oligocene of France as revealed by micro-computed tomography. Journal of Vertebrate Paleontology,37, e1384738.

    Google Scholar 

  69. Čerňanský, A., Daza, J. D., & Bauer, A. M. (2018). Geckos from the middle Miocene of Devínska Nová Ves (Slovakia): new material and a review of the previous record. Swiss Journal of Geosciences,111, 183–190.

    Google Scholar 

  70. Čerňanský, A., Klembara, J., & Müller, J. (2016a). The new rare record of the late Oligocene lizards and amphisbaenians from Germany and its impact on our knowledge of the European terminal Palaeogene. Palaeobiodiversity and Palaeoenvironments,96, 559–587.

    Google Scholar 

  71. Čerňanský, A., Klembara, J., & Smith, K. T. (2016b). Fossil lizard from central Europe resolves the origin of large body size and herbivory in giant Canary Island lacertids. Zoological Journal of the Linnean Society,176, 861–877.

    Google Scholar 

  72. Čerňanský, A., Rage, J.-C., & Klembara, J. (2015). The Early Miocene squamates of Amöneburg (Germany): the first stages of modern squamates in Europe. Journal of Systematic Palaeontology,13, 97–128.

    Google Scholar 

  73. Čerňanský, A., & Smith, K. T. (2018). Eolacertidae: a new extinct clade of lizards from the Palaeogene; with comments on the origin of the dominant European reptile group – Lacertidae. Historical Biology,30, 994–1014.

    Google Scholar 

  74. Čerňanský, A., Szyndlar, Z., & Mörs, T. (2017b). Fossil squamate faunas from the Neogene of Hambach (northwestern Germany). Palaeobiodiversity and Palaeoenvironments,97, 329–354.

    Google Scholar 

  75. Čerňanský, A., & Venczel, M. (2011). An amphisbaenid reptile (Squamata, Amphisbaenidae) from the lower Miocene of Northwest Bohemia (MN 3, Czech Republic). Neues Jahrbuch für Geologie und Paläontologie-Abhandlungen,260, 73–77.

    Google Scholar 

  76. Cirilli, O., Benvenuti, M. G., Carnevale, G., Casanovas Vilar, I., Delfino, M., Furió, M., et al. (2016). Fosso della Fittaia: the oldest Tusco-Sardinian late Miocene endemic vertebrate assemblages (Baccinello-Cinigiano Basin, Tuscany, Italy). Rivista Italiana di Paleontologia e Stratigrafia,122, 13–34.

    Google Scholar 

  77. Colombero, S., Alba, D. M., D’amico, C., Delfino, M., Esu, D., Giuntelli, P., et al. (2017). Late Messinian mollusks and vertebrates from Moncucco Torinese, North-Western Italy. Paleoecological and paleoclimatological implications. Palaeontologia Electronica, 20.1.10A, 1–66.

  78. Colombero, S., Angelone, C., Bonelli, E., Carnevale, G., Cavallo, O., Delfino, M., et al. (2014). The upper Messinian assemblages of fossil vertebrate remains of Verduno (NW Italy): another brick for a latest Miocene bridge across the Mediterranean. Neues Jahrbuch für Geologie und Paläontologie-Abhandlungen,272, 287–324.

    Google Scholar 

  79. Conrad, J. L. (2008). Phylogeny and systematics of Squamata (Reptilia) based on morphology. Bulletin of the American Museum of Natural History,310, 1–182.

    Google Scholar 

  80. Conrad, J. L., Ast, J. C., Montanari, S., & Norell, M. A. (2010). A combined evidence phylogenetic analysis of Anguimorpha (Reptilia: Squamata). Cladistics,16, 1–48.

    Google Scholar 

  81. Conrad, J. L., Balcarcel, A., & Mehling, C. (2012). Earliest example of a giant monitor lizard (Varanus, Varanidae, Squamata). PLoS ONE,7, e41767.

    Google Scholar 

  82. Crespo, E. G. (2001). Paleo-herpetofauna de Portugal. Lisbon: Museu Bocage.

    Google Scholar 

  83. Daza, J. D., Bauer, A. M., & Snively, E. D. (2014). On the fossil record of the Gekkota. The Anatomical Record,97, 433–462.

    Google Scholar 

  84. De Gregorio, A. (1925). Mammiferi quaternari di Sicilia (Microfauna). Mammiferi e altri vertebrati (Myoxus, Lepus, Lagomys, Pellegrinia, Mustela, Crocidura, Lacerta e Testudo etc.). Annales de Geologie et Paléontologie,43, 1–19.

    Google Scholar 

  85. de Rochebrune, A. (1884). Faune ophiologique des Phosphorites du Quercy. Mémoires de la Société des sciences naturelles de Saône-et-Loire,5, 149–164.

    Google Scholar 

  86. Delfino, M. (1997a). Italian paleoerpetofauna database: Neogene-Quaternary. In Z. Roček, & S. Hart (Eds.), Herpetology ‘97. Abstract of the III World Congress of Herpetology, 210 August 1997, Prague, Czech Republic (51–52.). Prague: Third World Congress of Herpetology.

  87. Delfino, M. (1997b). Blanus from the Early Pleistocene of Southern Italy: another small tessera from a big mosaic. In W. Böhme, W. Bischoff, & T. Ziegler (Eds.), Herpetologia Bonnensis (89–97.). Bonn: Societas Europaea Herpetologica.

  88. Delfino, M. (2002). Erpetofaune Italiane del Neogene e del Quaternario. Modena: University of Modena and Reggio Emilia.

    Google Scholar 

  89. Delfino, M. (2003). A Pleistocene amphisbaenian from Sicily. Amphibia-Reptilia,24, 407–414.

    Google Scholar 

  90. Delfino, M. (2004). The Middle Pleistocene herpetofauna of Valdemino Cave (Liguria, North-Western Italy). Herpetological Journal,14, 113–128.

    Google Scholar 

  91. Delfino, M. (2006). Il registro fossile della moderna erpetofauna italiana. In R. Sindaco, G. Doria, E. Razzetti, & F. Bernini (Eds.), Atlante degli anfibi e dei rettili d’Italia/Atlas of Italian amphibians and reptiles (pp. 96–119). Firenze: Societas Herpetologica Italica, Edizioni Polistampa.

    Google Scholar 

  92. Delfino, M. (2013). Cenozoic herpetofaunas of Apulia (Southern Italy). In G. Scillitani, C. Liuzzi, L. Lorusso, F. Mastropasqua, & P. Ventrella (Eds.), Atti IX Congresso Nazionale della Societas Herpetologica Italica (Bari - Conversano, 26–30 settembre 2012) (pp. 99–103). Conversano: Tipolitografia Pineta.

    Google Scholar 

  93. Delfino, M., & Bailon, S. (2000). Early Pleistocene herpetofauna from Cava Dell’Erba and Cava Pirro (Apulia, Southern Italy). Herpetological Journal,10, 95–110.

    Google Scholar 

  94. Delfino, M., Bailon, S., & Pitruzzella, G. (2011). The late Pliocene amphibians and reptiles from “Capo Mannu D1 Local Fauna” (Mandriola, Sardinia, Italy). Geodiversitas,33, 357–382.

    Google Scholar 

  95. Delfino, M., Kotsakis, T., Arca, M., Tuveri, C., Pitruzzella, G., & Rook, L. (2008). Agamid lizards from the Plio-Pleistocene of Sardinia (Italy) and an overview of the European fossil record of the family. Geodiversitas,30, 641–656.

    Google Scholar 

  96. Delfino, M., Rage, J.-C., Bolet, A., & Alba, D. M. (2013). Early Miocene dispersal of the lizard Varanus into Europe: reassessment of vertebral material from Spain. Acta Palaeontologica Polonica,58, 731–735.

    Google Scholar 

  97. Delfino, M., & Rook, L. (2008). The fossil amphibians and reptiles of Sardinia: a summary. In C. Corti (Ed.), Herpetologia Sardiniae (pp. 192–195). Latina: Societas Herpetologica Italica/Edizioni Belvedere.

    Google Scholar 

  98. Depéret, C. (1890). Les animaux pliocènes du Roussillon. Mémoires de la Société géologique de France, Paléontologie,3, 1–194.

    Google Scholar 

  99. Estes, R. (1969). Die Fauna der Miozänen Spaltenfüllung von Neudorf an der March (CSSR). Reptilia (Lacertilia). Sitzungsberichte der Akademie der Wissenschaften mathematisch-naturwissenschaftliche Klasse, 178, 77–82.

  100. Estes, R. (1983). Handbuch der Paläoherpetologie 10A. Sauria terrestria, Amphisbaenia. Munich: Friedrich Pfeil.

  101. Evans, E. S. (1991). A new lizard-like reptile (Diapsida: Lepidosauromorpha) from the Middle Jurassic of England. Zoological Journal of the Linnean Society,103, 391–412.

    Google Scholar 

  102. Evans, S. E. (1994). A new anguimorph lizard from the Jurassic and lower Cretaceous of England. Palaeontology,37, 33–49.

    Google Scholar 

  103. Evans, S. E. (1998). Crown group lizards (Reptilia, Squamata) from the Middle Jurassic of the British Isles. Palaeontographica, Abteilung A,250, 123–154.

    Google Scholar 

  104. Evans, S. E. (2008). The skull of lizards and Tuatara. In C. Gans, A. Gaunt., & K. Adler (Eds.), Biology of the Reptilia, vol. 20 (The skull of Lepidosauria). Contributions to Herpetology, 23, 1–347.

  105. Evans, S. E., Barrett, P. M., Hilton, J., Butler, R., Jones, M. E. H., Liang, M. M., et al. (2006). The Middle Jurassic vertebrate assemblage of Skye, Scotland. In S. E. Evans, & P. M. Barrett (Eds.), Ninth Symposium on Mesozoic Terrestrial Ecosystems and Biota (36–39.). London: Natural History Museum Publications.

  106. Evans, S. E., & Milner, A. R. (1994). Middle Jurassic microvertebrate assemblages from the British Isles. In N. C. Fraser & E. D. Sues (Eds.), In the shadow of the dinosaurs: Early Mesozoic tetrapods (pp. 303–321). Cambridge: Cambridge University Press.

    Google Scholar 

  107. Evans, S. E., Prasad, G. V. R., & Manhas, B. K. (2002). Fossil lizards from the Jurassic Kota Formation of India. Journal of Vertebrate Paleontology,22, 299–312.

    Google Scholar 

  108. Ferrandini, J., Salotti, M., Bailon, S., Bonifay, M. F., Mourer-Chauviré, C., & Real-Testud, A. M. (1995). Découverte d’importants remplissages fossilifères d’âge pléistocène supérieur et holocène dans le karst de la région d’Oletta (Haute Corse). Geobios,28, 117–124.

    Google Scholar 

  109. Folie, A., Smith, R., & Smith, T. (2013). New amphisbaenian lizards from the Early Paleogene of Europe and their implications for the early evolution of modern amphisbaenians. Geologica Belgica,16, 227–235.

    Google Scholar 

  110. Fraser, N. C. (1982). A new rhynchocephalian from the British Upper Trias. Palaeontology,25, 709–725.

    Google Scholar 

  111. Fraser, N. C., & Benton, M. J. (1989). The Triassic reptiles Brachyrhinodon and Polysphenodon and the relationships of the sphenodontids. Zoological Journal of the Linnean Society,96, 413–445.

    Google Scholar 

  112. Gamble, T., Greenbaum, E., Jackman, T. R., Russell, A. P., & Bauer, A. M. (2012). Repeated origin and loss of adhesive toepads in geckos. PLoS One,7, e39429.

    Google Scholar 

  113. Gamble, T., Greenbaum, E., Jackman, T. R., Russell, A. P., & Bauer, A. M. (2015). Into the light: diurnality has evolved multiple times in geckos. Biological Journal of the Linnean Society,115, 896–910.

    Google Scholar 

  114. García-Porta, J., Quintana, J., & Bailon, S. (2002). Primer hallazgo de Blanus sp. (Amphisbaenia, Reptilia) en el Neógeno balear. Revista Española de Herpetología,16, 19–28.

    Google Scholar 

  115. Gaudry, A. (1862). Animaux fossiles et géologie de l’Attique d’après les recherchers faites en 1855–56 et an 1860. Paris: F. Savy Editeur.

    Google Scholar 

  116. Gauthier, J., Kearney, M., Maisano, J. A., Rieppel, O., & Behlke, A. D. B. (2012). Assembling the squamate tree of life: perspectives from the phenotype and the fossil record. Bulletin of the Peabody Museum of Natural History,53, 3–308.

    Google Scholar 

  117. Georgalis, G. L. (2017). Necrosaurus or Palaeovaranus? Appropriate nomenclature and taxonomic content of an enigmatic fossil lizard clade (Squamata). Annales de Paléontologie,103, 293–303.

    Google Scholar 

  118. Georgalis, G. L., Villa, A., & Delfino, M. (2016a). First description of a fossil chamaeleonid from Greece and its relevance for the European biogeographic history of the group. The Science of Nature,103, 1–12.

    Google Scholar 

  119. Georgalis, G. L., Villa, A., & Delfino, M. (2017a). Fossil lizards and snakes from Ano Metochi—a diverse squamate fauna from the latest Miocene of northern Greece. Historical Biology,29, 730–742.

    Google Scholar 

  120. Georgalis, G. L., Villa, A., & Delfino, M. (2017b). The last European varanid: demise and extinction of monitor lizards (Squamata, Varanidae) from Europe. Journal of Vertebrate Paleontology,37, e1301946.

    Google Scholar 

  121. Georgalis, G. L., Villa, A., & Delfino, M. (2018). The last amphisbaenian (Squamata) from continental Eastern Europe. Annales de Paléontologie,104, 155–159.

    Google Scholar 

  122. Georgalis, G. L., Villa, A., Vlachos, E., & Delfino, M. (2016b). Fossil amphibians and reptiles from Plakias, Crete: a glimpse into the earliest late Miocene herpetofaunas of southeastern Europe. Geobios,49, 433–444.

    Google Scholar 

  123. Gerhardt, K. (1903). Ophisaurus ulmensis n. sp. aus dem Untermiozän von Ulm a. D. Jahreshefte des Vereins für Vaterländische Naturkunde in Württemberg,59, 67–71.

    Google Scholar 

  124. Gervais, P. (1859). Zoologie et Paléontologie Françaises. Paris: Arthus Bertrand.

    Google Scholar 

  125. Gulia, G. (1912). La geologia ed i fossili delle Isole Maltesi. In G. Muscat (Ed.), General Guide to Malta and Gozo for the year 1912 (pp. 291–318). Valletta: The Malta Herald.

    Google Scholar 

  126. Gulia, G. (1914). Uno sguardo alla zoologia delle “Isole Maltesi”. In L. Jourin (Ed.), IX congrès international de zoologie tenu á Monaco du 25 au 30 mars 1913 (pp. 545–555). Imprimerie Oberthür: Rennes.

    Google Scholar 

  127. Gvoždík, V., Benkovský, N., Crottini, A., Bellati, A., Moravec, J., Romano, A., et al. (2013). An ancient lineage of slow worms, genus Anguis (Squamata: Anguidae), survived in the Italian Peninsula. Molecular Phylogenetics and Evolution,69, 1077–1092.

    Google Scholar 

  128. Gvoždík, V., Jandzik, D., Lymberakis, P., Jablonski, D., & Moravec, J. (2010). Slow worm, Anguis fragilis (Reptilia: Anguidae) as a species complex: Genetic structure reveals deep divergences. Molecular Phylogenetics and Evolution,55, 460–472.

    Google Scholar 

  129. Harris, D. J., Batista, V., Carretero, M. A., & Ferrand, N. (2004). Genetic variation in Tarentola mauritanica (Reptilia: Gekkonidae) across the Strait of Gibraltar derived from mitochondrial and nuclear DNA sequences. Amphibia-Reptilia,25, 451–459.

    Google Scholar 

  130. Hedges, S. B., & Vidal, N. (2009). Lizards, snakes, and amphisbaenians (Squamata). In S. B. Hedges & S. Kumar (Eds.), The timetree of life (pp. 383–389). New York: Oxford University Press.

    Google Scholar 

  131. Hoffstetter, R. (1942). Sur la présence d’Amphisbaenidae dans les gisements tertiaires français. Comptes rendues des Séances de la Societé de Géologie de France,3–4, 24–25.

    Google Scholar 

  132. Hoffstetter, R. (1943). Varanidae et Necrosauridae fossiles. Bulletin du Muséum National d’Histoire Naturelle,15, 134–141.

    Google Scholar 

  133. Hoffstetter, R. (1944). Sur les Scincidae fossiles. I. Formes européennes et nord-américaines. Bulletin du Muséum National d’Histoire Naturelle, Paris,16, 547–553.

    Google Scholar 

  134. Hoffstetter, R. (1946). Sur les Gekkonidae fossiles. Bulletin du Muséum National d’Histoire Naturelle,18, 195–203.

    Google Scholar 

  135. Hoffstetter, R. (1969). Présence de Varanidae (Reptilia, Sauria) dans le Miocène de Catalogne. Considérations sur l’histoire de la famille. Bulletin du Muséum National d’Histoire Naturelle,40, 1051–1064.

    Google Scholar 

  136. Holman, J. A. (1998). Pleistocene amphibians and reptiles in Britain and Europe. Oxford monographs on geology and geophysics,38, 1–254.

    Google Scholar 

  137. Holmes, R. B., Murray, A. M., Attia, Y. S., Simons, E. L., & Chatrath, P. (2010). Oldest known Varanus (Squamata: Varanidae) from the upper Eocene and lower Oligocene of Egypt: support for an African origin of the genus. Palaeontology,53, 1099–1110.

    Google Scholar 

  138. Ivanov, M., Ruta, M., Klembara, J., & Böhme, M. (2018). A new species of Varanus (Anguimorpha: Varanidae) from the early Miocene of the Czech Republic, and its relationships and palaeoecology. Journal of Systematic Palaeontology,16, 767–797.

    Google Scholar 

  139. Jánossy, D. (1986). Pleistocene vertebrate faunas of Hungary. Budapest: Elsevier/Akadémiai Kiadó.

    Google Scholar 

  140. Jones, M. E. H., Anderson, C. L., Hipsley, C. A., Müller, J., Evans, S. E., & Schoch, R. R. (2013). Integration of molecules and new fossils supports a Triassic origin for Lepidosauria (lizards, snakes, and tuatara). BMC Evolutionary Biology,13, 208.

    Google Scholar 

  141. Jörg, E. (1965). Ophisaurus acuminatus nov. spec. (Anguidae, Rept.) von der pontischen Wirbeltier-Fundstätte Höwenegg Hegau. Beiträge zur naturkundlichen Forschungen in SW-Deutschland,24, 21–30.

    Google Scholar 

  142. Jost, J., Kälin, D., Börner, S., Vasilyan, D., Lawver, D., & Reichenbacher, B. (2015). Vertebrate microfossils from the Upper Freshwater Molasse in the Swiss Molasse Basin: implications for the evolution of the North Alpine Foreland Basin during the Miocene Climate Optimum. Palaeogeography, Palaeoclimatology, Palaeoecology,426, 22–33.

    Google Scholar 

  143. Jovanović, M., Đurić, D., & Marković, Z. (2002). Tertiary reptiles of the central part of the Balkan peninsula. Biota,3, 67–75.

    Google Scholar 

  144. Karin, B. R., Metallinou, M., Weinell, J. L., Jackman, T. R., & Bauer, A. M. (2016). Resolving the higher-order phylogenetic relationships of the circumtropical Mabuya group (Squamata: Scincidae): an out-of-Asia diversification. Molecular Phylogenetics and Evolution,102, 220–232.

    Google Scholar 

  145. Kearney, M. (2003). Systematics of the Amphisbaenia (Lepidosauria: Squamata) based on morphological evidence from recent and fossil forms. Herpetological Monographs,17, 1–74.

    Google Scholar 

  146. Klembara, J. (1979). Neue funde der gattungen Ophisaurus und Anguis (Squamata, Reptilia) aus dem Untermiozän Westböhmens (ČSSR). Vestník Ústředního Ústavu Geologického,54, 163–170.

    Google Scholar 

  147. Klembara, J. (1986). Neue funde der gattungen Pseudopus und Anguis (Reptilia, Anguinae) aus dei Pliopleistozänen Mitteleuropäischen lokalitäten. Geologica Carpathica,37, 91–106.

    Google Scholar 

  148. Klembara, J. (2008). A new anguimorph lizard from the lower Miocene of North-West Bohemia, Czech Republic. Palaeontology,51, 81–94.

    Google Scholar 

  149. Klembara, J. (2015). New finds of anguines (Squamata, Anguidae) from the early Miocene of Northwest Bohemia (Czech Republic). Paläontologische Zeitschrift,89, 171–195.

    Google Scholar 

  150. Klembara, J., Böhme, M., & Rummel, M. (2010). Revision of the anguine lizard Pseudopus laurillardi (Squamata, Anguidae) from the Miocene of Europe, with comments on paleoecology. Journal of Paleontology,84, 159–196.

    Google Scholar 

  151. Klembara, J., & Green, B. (2010). Anguimorph lizards (Squamata, Anguimorpha) from the middle and late Eocene of the Hampshire Basin of southern England. Journal of Systematic Palaeontology,8, 97–129.

    Google Scholar 

  152. Klembara, J., Hain, M., & Čerňanský, A. (2017). The first record of anguine lizards (Anguimorpha, Anguidae) from the early Miocene locality Ulm—Westtangente in Germany. Historical Biology, https://doi.org/10.1080/08912963.2017.1416469

    Google Scholar 

  153. Klembara, J., Hain, M., & Dobiašová, K. (2014). Comparative anatomy of the lower jaw and dentition of Pseudopus apodus and the interrelationships of species of subfamily Anguinae (Anguimorpha, Anguidae). The Anatomical Record,297, 516–544.

    Google Scholar 

  154. Klembara, J., & Rummel, M. (2018). New material of Ophisaurus, Anguis and Pseudopus (Squamata, Anguidae, Anguinae) from the Miocene of the Czech Republic and Germany and systematic revision and palaeobiogeography of the Cenozoic Anguinae. Geological Magazine,155, 20–44.

    Google Scholar 

  155. Kosma, R. (2004). The dentitions of recent and fossil scincomorphan lizards (Lacertilia, Squamata)—systematics, functional morphology, palecology. Hannover: Universität Hannover.

    Google Scholar 

  156. Kotsakis, T. (1977). I resti di anfibi e rettili pleistocenici della grotta di Spinagallo (Siracusa, Sicilia). Geologica Romana,16, 211–229.

    Google Scholar 

  157. Kretzoi, M., & Poulianos, N. (1981). Remarks on the middle and lower Pleistocene vertebrate fauna in the Petralona Cave. Anthropos,8, 57–72.

    Google Scholar 

  158. Lartet, E. (1851). Notice sur la colline de Sansan. Auch: J.-A. Portes.

    Google Scholar 

  159. Lungu, A. N., Zerova, G. A., & Chkhikvadze, V. M. (1983). Pervie svedeniia o miotsenovom varane severnogo prichernomoriia. Soobshcheniya Akademii Nauk Gruziinskoi SSR,110, 417–420.

    Google Scholar 

  160. Mangili, G. (1980). Fossils reptiles of Simonelli cave. Quaderni della Accademia Nazionale dei Lincei,249, 121–122.

    Google Scholar 

  161. Mateo, J. A. (1988). Estudio sistemático y zoogeográfico de los lagartos ocelados, Lacerta lepida Daudin, 1802, y Lacerta pater (Lataste, 1880) (Sauria: Lacertidae). Sevilla: University of Sevilla.

    Google Scholar 

  162. Meszoely, C. A. M., & Gasparik, M. (2002). First record of an agamid lizard from the Pleistocene of Hungary. Fragmenta Palaeontologica Hungarica,20, 1–2.

    Google Scholar 

  163. Meszoely, C. A. M., Schaff, C. R., & Jenkins, F., Jr. (1987). Early Jurassic sphenodontians from northeast Arizona. Journal of Vertebrate Paleontology,7, 21A.

    Google Scholar 

  164. Miklas-Tempfer, P. M. (2003). The Miocene herpetofaunas of Grund (Caudata; Chelonii, Sauria, Serpentes) and Mülbach am Manhartsberg (Chelonii, Sauria, Amphisbaenia, Serpentes), Lower Austria. Annalen des Naturhistorischen Museums in Wien,104A, 195–235.

    Google Scholar 

  165. Młynarski, M. (1956). Lizards from the Pliocene of Poland. Acta Palaeontologica Polonica,1, 135–152.

    Google Scholar 

  166. Młynarski, M. (1962). Notes on the amphibian and reptilian fauna of the Polish Pliocene and Early Pleistocene. Acta Zoologica Cracoviensia,7, 177–194.

    Google Scholar 

  167. Montoya, P., Alberdi, M. T., Barbadillo, L. J., Van Der Made, J., Morales, J., Murelaga, X., et al. (2001). Une faune très diversifiée du Pléistocène inférieur de la Sierra de Quibas (province de Murcia, Espagne). Comptes Rendus de l’Académie des Sciences, Earth and Planetary Science,332, 387–393.

    Google Scholar 

  168. Moody, S., & Roček, Z. (1980). Chamaeleo caroliquarti (Chamaeleonidae, Sauria) a new species from the lower Miocene of central Europe. Věstnik Ústředniho Ustavu Geologického,55, 85–92.

    Google Scholar 

  169. Morelli, N. (1891). Resti organici rinvenuti nella caverna delle Arene Candide. Atti della Società Ligustica di Scienze Naturali e Geografiche,2, 171–205.

    Google Scholar 

  170. Mukhopadhyay, G., Mukhopadhyay, S. K., Roychowdhury, M., & Parui, P. K. (2010). Stratigraphic correlation between different Gondwana basins of India. Journal of the Geological Society of India,76, 251–266.

    Google Scholar 

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

    Google Scholar 

  172. Müller, J. (2001). A new fossil species of Euleptes from the early Miocene of Montaigu, France (Reptilia, Gekkonidae). Amphibia-Reptilia,22, 341–348.

    Google Scholar 

  173. Müller, J., & Mödden, C. (2001). A fossil leaf-toed gecko from the Oppenheim-Nierstein Quarry (Lower Miocene, Germany). Journal of Herpetology,35, 529–532.

    Google Scholar 

  174. Nopcsa, F. (1908). Zur Kenntnis der fossilen Eidechsen. Beiträge zur Paläontologie von Österreich,21, 33–62.

    Google Scholar 

  175. Papp, A., Thenius, E., Berger, W., & Weinfurter, E. (1953). Vösendorf - ein Lebensbild aus dem Pannon des Wiener Beckens. Ein beitrag zur geologie und paläontologie der unterpliozänen congerienschichten des südlichen Wiener Beckens. Mitteilungen der Geologischen Gesellschaft in Wien,46, 1–109.

    Google Scholar 

  176. Pomel, M. (1853). Catalogue méthodique et descriptif des vertébrés fossiles découverts dans le bassin hydrogeographique supérieur de la Loire, et surtout dans la vallée de son affluent principal, l´Allier. Paris: J.-B. Baillière.

    Google Scholar 

  177. Prasad, G. V. R., & Manhas, B. K. (2007). A new docodont mammal from the Jurassic Kota Formation of India. Palaeontologia Electronica,10, 7A.

    Google Scholar 

  178. Pyron, R. A., Burbrink, F. T., & Wiens, J. J. (2013). A phylogeny and revised classification of Squamata, including 4161 species of lizards and snakes. BMC Evolutionary Biology,13, 93.

    Google Scholar 

  179. Rage, J.-C. (2013). Mesozoic and Cenozoic squamates of Europe. Palaeobiodiversity and Palaeoenvironments,93, 517–534.

    Google Scholar 

  180. Rage, J.-C., & Bailon, S. (2005). Amphibians and squamate reptiles from the late early Miocene (MN 4) of Béon 1 (Montréal-du-Gers, southwestern France). Geodiversitas,27, 413–441.

    Google Scholar 

  181. Rage, J.-C., & Szyndlar, Z. (2005). Latest Oligocene–early Miocene in Europe: dark Period for booid snakes. Comptes Rendus Palevol,4, 428–435.

    Google Scholar 

  182. Rato, C., Carranza, S., & Harris, D. J. (2011). When selection deceives phylogeographic interpretation: the case of the Mediterranean house gecko, Hemidactylus turcicus (Linnaeus, 1758). Molecular Phylogenetics and Evolution,58, 365–373.

    Google Scholar 

  183. Rato, C., Carranza, S., Perera, A., Carretero, M. A., & Harris, D. J. (2010). Conflicting patterns of nucleotide diversity between mtDNA and nDNA in the Moorish gecko, Tarentola mauritanica. Molecular Phylogenetics and Evolution,56, 962–971.

    Google Scholar 

  184. Rauscher, K. L. (1992). Die Echsen (Lacertilia, Reptilia) aus dem Plio-Pleistozän von Bad Deutsch-Altenburg, Niederösterreich. Beiträge zur paläontologie von Österreich,17, 81–177.

    Google Scholar 

  185. Rauscher, K. L. (1995). Die herpetofauna der Vraona-Hölhe (Attika) in Griechenland. Annales Géologiques des Pays Helléniques,36, 39–41.

    Google Scholar 

  186. Renesto, S., & Posenato, R. (2003). A new lepidosauromorph reptile from the Middle Triassic of the Dolomites (Northern Italy). Rivista Italiana di Paleontologia e Stratigrafia,109, 463–474.

    Google Scholar 

  187. Reynoso, V. H. (1998). Huehuecuetzpali mixtecus gen. et sp. nov: a basal squamate (Reptilia) from the Early Cretaceous of Tepexi de Rodríguez, Central México. Philosophical Transactions of the Royal Society of London B, Biological Sciences,353, 477–500.

    Google Scholar 

  188. Roček, Z. (1984). Lizards (Reptilia: Sauria) from the lower Miocene locality Dolnice (Bohemia, Czechoslovakia). Rozpravy Československé Akademie Věd - Řada Matematických a Přírodních Věd,94, 3–64.

    Google Scholar 

  189. Roger, O. (1898). Wirbelthierreste aus dem Dinotheriensande, II. Theil. Bericht des Naturwissenschaftlichen Vereins fur Schwaben und Neuburg (a.V.) in Augsburg, 33, 385–396.

  190. Rook, L., Abbazzi, L., Angelone, C., Arca, M., Barisone, G., Bedetti, C., et al. (2003). Osservazioni preliminari sui vertebrati fossili plio-pleistocenici del Monte Tuttavista (Orosei, Sardegna). International Journal of Archaeology—Sardinia Corsica et Baleares Antiquae,1, 11–29.

    Google Scholar 

  191. Salotti, M., Bailon, S., Bonifay, M.-F., Courtois, J.-Y., Dubois, J.-N., Ferrandini, J., et al. (1997). Castiglione 3, un nouveau remplissage fossilifère d’âge Pléistocène moyen dans le karst de la région d’Oletta (Haute-Corse). Comptes Rendus de l’Académie des Sciences, Paris, IIa,324, 67–74.

    Google Scholar 

  192. Salotti, M., Louchart, A., Bailon, S., Lorenzo, S., Oberlin, C., et al. (2008). A Teppa di U Lupinu Cave (Corsica, France)—human presence since 8500 years BC, and the enigmatic origin of the earlier, late Pleistocene accumulation. Acta Zoologica Cracoviensia,51A, 15–34.

    Google Scholar 

  193. Sampaio, F. L., Harris, D. J., Perera, A., & Salvi, D. (2015). Phylogenetic and diversity patterns of Blanus worm lizards (Squamata: Amphisbaenia): insights from mitochondrial and nuclear gene genealogies and species tree. Journal of Zoological Systematics and Evolutionary Research,53, 45–54.

    Google Scholar 

  194. Savona Ventura, C. (1984). The fossil herpetofauna of the Maltese islands, a review. Naturalista Siciliano,8, 93–106.

    Google Scholar 

  195. Schleich, H. H. (1983). Die mittelmiozäne Fossil-Lagerstätte Sandelzhausen. 13. Chamaeleo bavaricus sp. nov., ein neuer Nachweis aus dem Jungtertiär Süddeutschlands. Mitteilungen der Bayerischen Staatssammlung für Paläontologie und Historische Geologie,23, 77–82.

    Google Scholar 

  196. Schleich, H. H. (1984). Neue Reptilienfunde aus dem Tertiär Deutschlands. 2. Chamaeleo pfeili sp. nov. von der untermiozänen Fossilfundstelle Rauscheröd/Niederbayern (Reptilia, Sauria, Chamaeleonidae). Mitteilungen der Bayerischen Staatssammlung für Paläontologie und Historische Geologie,24, 97–104.

    Google Scholar 

  197. Schleich, H. H. (1985). Neue reptilienfunde aus dem Tertiär Deutschlands. 3. Erstnachweis von Doppelschleichen (Blanus antiquus sp.nov.) aus dem Mittelmiozän Süddeutschlands. Münchner Geowissenschaftliche Abhandlungen Reihe A,4, 1–16.

    Google Scholar 

  198. Schleich, H. H. (1987). Neue reptilienfunde aus dem Tertiär Deutschlands. 7. Erstnachweis von Geckos aus dem Mittelmiozän Süddeutschlands: Palaeogekko risgoviensis nov. gen., nov. spec. (Reptilia, Sauria, Gekkonidae). Mitteilungen der Bayerischen Staatssammlung für Palaeontologie und Historische Geologie,27, 67–93.

    Google Scholar 

  199. Schleich, H. H. (1988). Neue reptilienfunde aus dem Tertiär Deutschlands 8. Palaeoblanus tobieni n.gen., n.sp.—Neue Doppelschleichen aus dem Tertiär Deutschlands. Paläontologische Zeitschrift,62, 95–105.

    Google Scholar 

  200. Schleich, H. H. (1994). Neue Reptilienfunde aus dem Tertiär Deutschlands 15. Neue Funde fossiler Chamäleonen aus dem Neogen Süddeutschlands. Courier Forschungsinstitut Senckenberg,173, 175–195.

    Google Scholar 

  201. Schmitz, A., Mausfeld, P., & Embert, D. (2004). Molecular studies on the genus Eumeces Wiegmann, 1834: phylogenetic relationships and taxonomic implications. Hamadryad,28, 73–89.

    Google Scholar 

  202. Sears, M. W., & Angilletta, M. J., Jr. (2004). Body size clines in Sceloporus lizards: proximate mechanisms and demographic constraints. Integrative and Comparative Biology,44, 433–442.

    Google Scholar 

  203. Sickenberg, O. (1971). Revision der wirbeltierfauna der Höhle Petralona (Griech, Mazedonien). Annales Géologiques des Pays Helléniques,23, 230–264.

    Google Scholar 

  204. Sillero, N., Campos, J., Bonardi, A., Corti, C., Creemers, R., Crochet, P.-A., et al. (2014). Updated distribution and biogeography of amphibians and reptiles of Europe. Amphibia-Reptilia,35, 1–31.

    Google Scholar 

  205. Simões, T. R., Caldwell, M. W., Tałanda, M., Bernardi, M., Palci, A., Vernygora, O., et al. (2018). The origin of squamates revealed by a Middle Triassic lizard from the Italian Alps. Nature,557, 706–709.

    Google Scholar 

  206. Sindaco, R., Kornilios, P., Sacchi, R., & Lymberakis, P. (2014). Taxonomic reassessment of Blanus strauchi (Bedriaga, 1884) (Squamata: Amphisbaenia: Blanidae), with the description of a new species from South-East Anatolia (Turkey). Zootaxa,3795, 311–326.

    Google Scholar 

  207. Smith, K. T. (2017). First crocodile-tailed lizard (Squamata: Pan-Shinisaurus) from the Paleogene of Europe. Journal of Vertebrate Paleontology,37, e1313743.

    Google Scholar 

  208. Smith, K. T., & Gauthier, J. A. (2013). Early Eocene lizards of the Wasatch Formation near Bitter Creek, Wyoming: diversity and paleoenvironment during an interval of global warming. Bulletin of the Peabody Museum of Natural History,54, 135–230.

    Google Scholar 

  209. Smith, K. T., Maul, L. C., Flemming, F., Barkai, R., & Gopher, A. (2016). The microvertebrates of Qesem Cave: a comparison of the two concentrations. Quaternary International,398, 233–245.

    Google Scholar 

  210. Smith, K. T., & Scanferla, A. (2016). Fossil snake preserving three trophic levels and evidence for an ontogenetic dietary shift. Palaeobiodiversity and Palaeoenvironments,96, 589–599.

    Google Scholar 

  211. Smith, K. T., Schaal, S. F. K., & Habersetzer, J. (2018). Messel: an ancient greenhouse ecosystem. Stuttgart: Schweizerbart.

    Google Scholar 

  212. Speybroeck, J., Beukema, W., Bok, B., & Van der Voort, J. (2016). Field guide to the amphibians and reptiles of Britain and Europe. London: Bloomsbury Publishing.

    Google Scholar 

  213. Swinton, W. E. (1939). A new Triassic rhynchocephalian from Gloucestershire. Annals and Magazine of Natural History: Zoology, Botany, and Geology,4, 591–594.

    Google Scholar 

  214. Talavera, R. R., & Sanchíz, B. (1983). Restos holocénicos de Camaleón común, Chamaeleo chamaeleon (L.) de Málaga. Boletín de la Real Sociedad Española de Historia Natural, Sección Geológica,81, 81–84.

    Google Scholar 

  215. Tempfer, P. M. (2003). Amphibians and reptiles of the Karpatian Central Paratethys. In R. Brzobohatý, I. Cicha, M. Kováč, & F. Rögl (Eds.), The Karpatian—a Lower Miocene Stage of the Central Paratethys (pp. 285–291). Brno: Masarykova Univerzita v Brně.

    Google Scholar 

  216. Tempfer, P. M. (2005). The herpetofauna (Amphibia: Caudata, Anura; Reptilia: Scleroglossa) of the upper Miocene locality Kohfidisch (Burgenland, Austria). Beiträge zur Paläontologie,29, 145–253.

    Google Scholar 

  217. Tschopp, E., Villa, A., Camaiti, M., Ferro, L., Tuveri, C., Rook, L., et al. (2018). The first fossils of Timon (Squamata: Lacertinae) from Sardinia (Italy) and potential causes for its local extinction in the Pleistocene. Zoological Journal of the Linnean Society, https://doi.org/10.1093/zoolinnean/zly003

    Google Scholar 

  218. Uetz, P., Freed, P., & Hošek, J. (Eds.). The Reptile Database, http://www.reptile-database.org, accessed April 2018.

  219. Venczel, M. (2006). Lizards from the late Miocene of Polgárdi (W-Hungary). Nymphaea: Folia Naturae Bihariae, 33, 25–38.

  220. Venczel, M., & Hír, J. (2013). Amphibians and squamates from the Miocene of Felsötárkány Basin, N-Hungary. Palaeontographica. Abt. A: Palaeozoology - Stratigraphy,300, 117–158.

    Google Scholar 

  221. Venczel, M., & Sanchiz, B. (2006). Lower Miocene amphibians and reptiles from Oschiri (Sardinia, Italy). Hantkeniana,5, 72–75.

    Google Scholar 

  222. Venczel, M., & Ştiucă, E. (2008). Late middle Miocene amphibians and squamate reptiles from Tauţ, Romania. Geodiversitas,30, 731–763.

    Google Scholar 

  223. Vigne, J.-D., Bailon, S., & Cuisin, J. (1997). Biostratigraphy of amphibians, reptiles, birds and mammals in Corsica and the role of man in the Holocene faunal turnover. Anthropozoologica,25(26), 587–604.

    Google Scholar 

  224. Vigne, J.-D., Bailon, S., Cuisin, J., & Desse-Berset, N. (2002). Sensibilité des microvertébrés aux fluctiations hygrométriques tardilaciaires et holocènes: deux séquences en zone méditerranéenne humide (Grítulu et Monte di Tuda, Haut-Corse). In J.-P. Bravard & M. Magny (Eds.), Histoire des rivières et des lacs de Lascaux à nos jours (pp. 269–277). Paris: Errance.

    Google Scholar 

  225. Vijaya, X., & Prasad, G. V. R. (2001). Age of the Kota formation, Pranhita-Godavari Valley, India: a palynological approach. Journal of the Palaeontological Society of India,46, 77–93.

    Google Scholar 

  226. Villa, A., Blain, H.-A., & Delfino, M. (2018a). The Early Pleistocene herpetofauna of Rivoli Veronese (Northern Italy) as evidence for humid and forested glacial phases in the Gelasian of Southern Alps. Palaeogeography, Palaeoclimatology, Palaeoecology,490, 393–403.

    Google Scholar 

  227. Villa, A., Blain, H.-A., Van Den Hoek Ostende, L. W., & Delfino, M. (2018b). Fossil amphibians and reptiles from Tegelen (Province of Limburg) and the early Pleistocene palaeoclimate of The Netherlands. Quaternary Science Reviews,187, 203–219.

    Google Scholar 

  228. Villa, A., Bon, M., & Delfino, M. (2018c). Trapped in a roman well: amphibians and reptiles from Tenuta Zuccarello near Marcon, Venice, Italy. Historical Biology. https://doi.org/10.1080/08912963.2018.1470170.

    Article  Google Scholar 

  229. Villa, A., Daza, J. D., Bauer, A. M., & Delfino, M. (2018d). Comparative cranial osteology of European gekkotans (Reptilia, Squamata). Zoological Journal of the Linnean Society. https://doi.org/10.1093/zoolinnean/zlx104.

    Article  Google Scholar 

  230. Villa, A., & Delfino, M. (2017). Southern Germany: an early to middle Miocene lizard melting pot? Zitteliana,91, 93.

    Google Scholar 

  231. Villa, A., Kirchner, M., Alba, D. M., Bernardini, F., Bolet, A., Luján, À. H., et al. (2017b). Comparative cranial osteology of extant and extinct Blanus (Squamata, Amphisbaenia). In M. Menegon, A. Rodriguez-Prieto, & M. C. Deflorian (Eds.), AttiXI Congresso Nazionale della Societas Herpetologica Italica. Trento, 2225 settembre 2016 (329–333). Pescara: Ianieri Edizioni.

  232. Villa, A., Kosma, R., Čerňanský, A., & Delfino, M. (2018e). Taxonomical assessment of ‘Bavaricordylus’ Kosma, 2004 (Reptilia, Squamata). Journal of Vertebrate Paleontology. https://doi.org/10.1080/02724634.2018.1487844.

    Article  Google Scholar 

  233. Villa, A., Rook, L., Sami, M., & Delfino, M. (2016). Amphibians and reptiles from Cava Monticino (Ravenna, Italy) in the frame of the late Miocene Italian herpetofaunas. In L. Rook, S. Bartolini, & E. Ghezzo (Eds.), Paleodays 2016. La Società Paleontologica Italiana a Faenza. XVI riunione annuale SPI, Faenza (82). Torino: Centro Stampa/Regione Piemonte.

  234. Villa, A., Tschopp, E., Georgalis, G. L., & Delfino, M. (2017a). Osteology, fossil record and palaeodiversity of the European lizards. Amphibia-Reptilia,38, 79–88.

    Google Scholar 

  235. Vitt, L. J., & Caldwell, J. P. (2009). Herpetology (3rd ed.). Burlington: Academic Press.

    Google Scholar 

  236. Weithofer, A. (1888). Beiträge zur Kenntniss der Fauna von Pikermi bei Athen. Beiträge Paläontologie Österreich-Ungarns,6, 225–292.

    Google Scholar 

  237. Whiteside, D. I. (1986). The head skeleton of the Rhaetian sphenodontid Diphydontosaurus avonis gen. et sp. nov. and the modernizing of a living fossil. Philosophical Transactions of the Royal Society of London. Series B, Biological Sciences,312, 379–430.

    Google Scholar 

  238. Zerova, G. A., & Chkhikvadze, V. M. (1986). Neogene varanids of the URSS. In Z. Rocek (Ed.), Studies in Herpetology (pp. 689–694). Prague: Societas Herpetologica Europaea.

    Google Scholar 

  239. Zittel, K. (1889). Handbuch der Paläontologie. Munich: R. Oldenbourg.

    Google Scholar 

Download references

Acknowledgements

This overview was part of the Ph.D. thesis of one of us (A.V.) at the University of Torino. It greatly benefited from discussions with Aaron Bauer (Villanova University), Arnau Bolet (University of Bristol), Georgios Georgalis (University of Fribourg), Emanuel Tschopp (American Museum of Natural History, New York City) and Davit Vasilyan (JURASSICA Museum, Porrentruy). Two reviewers, Krister Smith (Senckenberg Research Institute, Frankfurt am Main) and Andrej Čerňanský (Comenius University, Bratislava), are thanked for useful comments on a previous version of this article. We would also like to thank the editor Daniel Marty. A.V. thanks Daniele Arobba, Andrea De Pascale and all the staff at Museo Archeologico del Finale (Finale Ligure) for their assistance while studying Ligurian fossil lizards, including the strange Hemidactylus from Valdemino. Oliver Rauhut (Bayerische Staatssammlung für Paläontologie und Geologie, Munich), Caterinella Tuveri (Soprintendenza Archeologia, Belle Arti e Paesaggio per le prov. di Sassari e Nuoro, Nuoro), Alexander Kupfer and Erin Maxwell (Staatliches Museum für Naturkunde, Stuttgart) kindly gave access to the collections under their care and helped while studying them. Visits to the Bayerische Staatssammlung für Paläontologie und Geologie (Munich) were supported by an EAVP Research Grant from the European Association of Vertebrate Palaeontologists to A.V. Arnau Bolet and David M. Alba (Institut Català de Paleontologia Miquel Crusafont, Sabadell) kindly allowed us to use their photos and the 3D model of B. mendezi in this paper, whereas Hugues-Alexandre Blain (Institut Català de Paleoecologia Humana i Evolució Social, Tarragona) gave us permission to figure his drawings of the Iberian Chalcides dentary represented in Fig. 8a–b. Project supported by Fondi di Ateneo (2016–2017), Generalitat de Catalunya (2014 SGR 416 GRC and CERCA Program), and Spanish Ministerio de Economía y Competitividad (CGL2016-76431-P). We would also like to acknowledge the Google Books team for having digitalized some classic palaeontological books, difficult to be accessed otherwise.

Author information

Affiliations

Authors

Corresponding author

Correspondence to Andrea Villa.

Additional information

Editorial Handling: M. R. Sánchez-Villagra.

Electronic supplementary material

Rights and permissions

Reprints and Permissions

About this article

Verify currency and authenticity via CrossMark

Cite this article

Villa, A., Delfino, M. Fossil lizards and worm lizards (Reptilia, Squamata) from the Neogene and Quaternary of Europe: an overview. Swiss J Palaeontol 138, 177–211 (2019). https://doi.org/10.1007/s13358-018-0172-y

Download citation

Keywords

  • “Lacertilia”
  • Amphisbaenia
  • fossil record
  • Cenozoic