Palaeobiodiversity and Palaeoenvironments

, Volume 96, Issue 1, pp 111–124 | Cite as

First record of Pelobates syriacus (Anura, Amphibia) in the early Pleistocene of Italy

  • Hugues-Alexandre Blain
  • Massimo Delfino
  • Claudio Berto
  • Marta Arzarello
Original Paper


Spadefoot toads (Pelobatidae, Anura) are rare in the Italian fossil record, being previously recorded only in the late Pliocene of Arondelli (Piedmont Region, northwestern Italy) and in the early Pleistocene of Pirro Nord P21 (Apulia, southeastern Italy). Here we describe for the first time abundant fossil remains of Pelobates syriacus from the lower Pleistocene fissure of Pirro Nord P13 (Apulia, southeastern Italy). These fossils shed new light on the palaeobiogeographical history of Pelobates in the Apennine Peninsula and are the first fossil evidence in Italy for P. syriacus, which currently lives in the southeastern Balkan Peninsula, Caucasus and the Middle East. This extralimital Italian occurrence suggests that P. syriacus has good dispersal abilities, when favourable conditions exist. Dispersal routes to reach the Apennine Peninsula during the early Pleistocene may have been favoured by a low level of the Adriatic Sea during a cold interval, furnishing a new habitat suitable for P. syriacus.


Pelobates Palaeobiogeography Trans-Adriatic distribution Early Pleistocene Italy 



We thank Jim Gardner (Drumheller, Canada) and Tomáš Přikryl (Prague, Czech Republic) for their invitation to participate in such a special issue in honour of Prof. Zbyněk Roček and Editor Sinje Weber for handling the manuscript. We are grateful to Prof. Zbyněk Roček (Department of Palaeobiology, Geological Institute, Prague) and Dr. Marta Calvo Revuelta (Museo Nacional de Ciencias Naturales, Madrid) for allowing us to study comparative specimens under their care. Tomáš Přikryl (Prague, Czech Republic), Márton Venczel (Oradrea, Romania), Vladimir Yuri Ratnikov (Voronezh, Russia), Rebecca Biton (Jerusalem, Israel), Ruben Iosif (Constanţa, Romania), Juan Manuel López-García (Tarragona, Spain), and Emilia Cianci (Torino, Italy) provided useful comments, references, or both. Jean-Claude Rage (Paris, France) and an anonymous reviewer improved the quality of the manuscript. Fieldwork at Pirro Nord P13 fissure was financially supported by Università degli Studi di Ferrara and Apricena Municipality. The excavations were possible thanks to the concession of the “Soprintendenza Archeologica della Puglia” and the “Ministero dei Beni e delle Attività Culturali”, to the collaboration of Dott. Ssa Anna Maria Tunzi, and to the kind disposability of Gaetano and Franco dell’Erba. This manuscript is part of projects CGL2011-13293-E/BTE, CGL2012-38358, SGR2014-901, 2014 SGR 416 GRC, and supported by Fondi di Ateneo dell’Università di Torino (2013-2014).


  1. Abbazzi, L., Benvenuti, M., Boschian, G., Dominici, S., Masini, F., Mezzabotta, C., Piccini, L., Rook, L., Valleri, G., & Torre, D. (1996). Revision of the Neogene and Pleistocene of the Gargano region (Apulia, Italy). The marine and continental successions and the mammal faunal assemblages in an area between Apricena and Poggio Imperiale (Foggia). Memorie Società Geologica Italiana, 51, 383–402.Google Scholar
  2. Alba, D. M., Colombero, S., Delfino, M., Martínez-Navarro, B., Pavia, M., & Rook, L. (2014). A thorny question: The taxonomic identity of the Pirro Nord cervical vertebrae revisited. Journal of Human Evolution, 76, 92–106.CrossRefGoogle Scholar
  3. Anastasakis, G., Piper, D. J. W., Dermitzakis, M. D., & Karakitsios, V. (2006). Upper Cenozoic stratigraphy and palaeogeographic evolution of Myrtoon and adjacent basins. Aegean Sea, Greece. Marine and Petroleum Geology, 23, 353–369.CrossRefGoogle Scholar
  4. Araújo, M. B., Thuiller, W., & Pearson, R. G. (2006). Climate warming and the decline of amphibians and reptiles in Europe. Journal of Biogeography, 33, 1712–1728.CrossRefGoogle Scholar
  5. Arzarello, M., & Peretto, C. (2010). Out of Africa: the first evidence of Italian peninsula occupation. Quaternary International, 223–224, 65–70.CrossRefGoogle Scholar
  6. Arzarello, M., Marcolini, F., Pavia, G., Pavia, M., Petronio, C., Petrucci, M., Rook, L., & Sardella, R. (2007). Evidence of earliest human occurrence in Europe: the site of Pirro Nord (southern Italy). Naturwissenschaften, 94, 107–112.CrossRefGoogle Scholar
  7. Arzarello, M., Marcolini, F., Pavia, G., Pavia, M., Petronio, C., Petrucci, M., Rook, L., & Sardella, R. (2009). L’industrie lithique du site Pléistocène inférieur de Pirro Nord (Apricena, Italie du Sud): une occupation humaine entre 1,3 et 1,7 Ma. L'Anthropologie, 113, 47–58.CrossRefGoogle Scholar
  8. Arzarello, M., Peretto, C., & Moncel, M.-H. (2015). The Pirro Nord site (Apricena, Fg, Southern Italy) in the context of the first European peopling: Convergences and divergentes. Quaternary International. doi: 10.1016/j.quaint.2014.08.051.Google Scholar
  9. Bailon, S. (1988). Amphibiens et reptiles. Rapport d'étude. In P. Van Ossel (Ed.), Les jardins du Carrousel (Paris). De la campagne à la ville: la formation d'un espace urbain. Documents d'Archéologie Française, 73, 277–283.Google Scholar
  10. Bailon, S. (1991). Amphibiens et reptiles du Pliocène et du Quaternaire de France et d’Espagne: mise en place et évolution des faunes. Unpublished PhD thesis. Université de Paris VII, Paris.Google Scholar
  11. Bailon, S. (1999a). Différenciation ostéologique des anoures (Amphibia, Anura) de France. In J. Desse & N. Desse-Berset (Eds.), Fiches d’ostéologie animale pour l’Archéologie, Série C Varia. Valbonne: Centre de Recherches Archéologiques—CNRS.Google Scholar
  12. Bailon, S. (1999b). Les amphibiens et les reptiles. In A. Defleur (Ed.), Rapport de synthèse de fouilles pluriannuelles, 1996–1999, Baume Moula-Guercy, Soyons, Ardèche, report for the Direction régionale des affaires culturelles (DRAC) Rhône-Alpes (pp. 111–127). Unpublished report.Google Scholar
  13. Bar-Yosef, O., & Tchernov, E. (1966). Archaeological finds and the fossil faunas of the Natufian and Microlithics industries at Hayonim cave, western Galilee, Israel. Israel Journal of Zoology, 15, 104–140.Google Scholar
  14. Bedetti, C. (2003). Le avifaune fossili del Plio-Pleistocene italiano: sistematica, paleoecologia ed elementi di biocronologia. Unpublished PhD thesis. Universita “La Sapienza” di Roma, Rome.Google Scholar
  15. Bermúdez de Castro, J. M., & Martinón-Torres, M. (2013). A new model for the evolution of the human Pleistocene populations of Europe. Quaternary International, 295, 102–112.CrossRefGoogle Scholar
  16. Bertini, A., Ciaranfi, N., Marino, M., & Palombo, M. R. (2010). Proposal for Pliocene and Pleistocene land-sea correlation in the Italian area. Quaternary International, 219, 95–108.CrossRefGoogle Scholar
  17. Blain, H.-A. (2014). Anfibios y escamosos de Cueva Victoria. In L. Gibert & C. Ferràndez-Cañadell (Eds.), Paleontología y Geología de Cueva Victoria, Cartagena, España. Mástia, 10, 161–183.Google Scholar
  18. Blain, H.-A., & Bailon, S. (2003). Les amphibiens et les reptiles des couches du Pléistocène supérieur ancien du gisement d’Artenac (Charente, France). Quaternaire, 14(2), 85–95.CrossRefGoogle Scholar
  19. Blain, H.-A., & Villa, P. (2006). Amphibians and squamate reptiles from the early upper Pleistocene of Bois Roche Cave (Charente, southwestern France). Acta Zoologica Cracoviensia, 49A(1-2), 1–32.CrossRefGoogle Scholar
  20. Blain, H.-A., Gibert, L., & Ferrandez-Cañadell, C. (2010). First report of a green toad (Bufo viridis sensu lato) in the early Pleistocene of Spain: Palaeobiogeographical and palaeoecological implications. Comptes Rendus Palevol, 9, 487–497.CrossRefGoogle Scholar
  21. Blain, H.-A., Bailon, S., & Agustí, J. (2015). The geographical and chronological pattern of herpetofaunal Pleistocene extinctions on the Iberian Peninsula. Comptes Rendus Palevol. doi: 10.1016/j.crpv.2015.05.008.Google Scholar
  22. Blaustein, A. R., Wake, D. B., & Sousa, W. P. (1994). Amphibian declines: judging stability, persistence, and susceptibility of populations to local and global extinctions. Conservation Biology, 8, 60–71.CrossRefGoogle Scholar
  23. 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 Zeitschift, 84, 3–41.CrossRefGoogle Scholar
  24. Böhme, M., & Ilg, A. (2003). fosFARbase. Available from: Accessed 18 Mar 2015.
  25. Böhme, W., Roček, Z., & Špinar, Z. V. (1982). On Pelobates decheni Troschel, 1861, and Zaphrissa eurypelis Cope, 1866 (Amphibia: Salientia: Pelobatidae) from the early Miocene of Rott near Bonn, West Germany. Journal of Vertebrate Paleontology, 2, 1–9.CrossRefGoogle Scholar
  26. Boettger, O. (1889). Ein neuer Pelobates aus Syrien. Zoologischer Anzeiger, 12, 144–147.Google Scholar
  27. Bologna, M. A., & Mazzotti, S. (2009). Biogeography. In R. Sindaco, G. Doria, E. Razzetti, & F. Bernini (Eds.), Atlas of Italian amphibians and reptiles (pp. 654–677). Edizioni Polistampa. Firenze: Societas Herpetologica Italica. Google Scholar
  28. Bonaparte, C. L. J. L. (1850). Conspectus Systematum. Herpetologiae et Amphibiologiae. Editio altera reformata. Lugdini Batavorum: E. J. Brill. Italica.Google Scholar
  29. Çıplak, B., Heller, K. G., & Willemse, F. (2010). Phylogeny and biogeography of Eupholidoptera Mařan (Orthoptera, Tettigoniidae): morphological speciation in correlation with the geographical evolution of the eastern Mediterranean. Systematic Entomology, 35, 722–738.CrossRefGoogle Scholar
  30. Delfino, M. (2002). Erpetofaune italiane del Neogene e del Quaternario. Unpublished PhD thesis. University of Modena and Reggio Emilia, Modena.Google Scholar
  31. Delfino, M., & Atzori, M. (2013). An update on the early Pleistocene herpetofauna from Pirro Nord. Palaeontographica Abteilung A, 298, 19–29.CrossRefGoogle Scholar
  32. 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
  33. Delfino, M., Bar-Oz, G., & Weissbrod, L. (2007). Recent shrinkage of the range of the Eastern Spadefoot Toad, Pelobates syriacus (Amphibia, Anura): archaeological evidence from the Bronze Age in Israel. Zoology in the Middle East, 40(1), 45–52.CrossRefGoogle Scholar
  34. Dermitzakis, M. D. (1990). Palaeogeography, geodynamic processes and event stratigraphy during the late Cenozoic of the Aegean area. International Symposium on Biogeographical Aspects of Insularities. Academia Nazionale Lincei, 85, 263–288.Google Scholar
  35. Džukić, G., Beškov, V., Sidorovska, V., Cogălniceanu, D., & Kalezić, L. M. (2005). Historical and contemporary ranges of the spadefoot toads Pelobates spp. (Amphibia: Anura) in the Balkan Peninsula. Acta Zoologica Cracoviensia, 48A(1-2), 1–9.Google Scholar
  36. Freudenthal, M. (1971). Neogene vertebrates from the Gargano Peninsula, Italy. Scripta Geologica, 3, 1–10.Google Scholar
  37. Frost, D. R. (2015) Amphibian Species of the Worlds: an Online Reference. Version 6.0 (1 April 2015). Electronic Database accessible at: New York: American Museum of Natural History.
  38. Geraga, M., Tsaila-Monopolis, S., Ioakim, C., Papatheodorou, G., & Ferentinos, G. (2000). Evaluation of palaeoenvironmental changes during the last 18,000 years in the Myrtoon Basin, SW Aegean Sea. Paleaeogeography, Palaeoclimatology, Palaeoecology, 156, 1–17.CrossRefGoogle Scholar
  39. Gleed-Owen, C. P. (1998). Quaternary herpetofaunas of the British Isles: taxonomic descriptions, palaeoenvironmental reconstructions, and biostratigraphic implications. Unpublished PhD thesis. Coventry University, Coventry.Google Scholar
  40. Gliozzi, E., Abbazzi, L., Argenti, P., Azzaroli, A., Caloi, L., Capasso Barbato, L., Di Stefano, G., Esu, D., Ficcarelli, G., Girotti, O., Kotsakis, T., Masini, F., Mazza, P., Mezzabotta, C., Palombo, M. R., Petronio, C., Rook, L., Sala, B., Sardella, R., Zanalda, E., & Torre, D. (1997). Biochronology of selected mammals, molluscs and ostracods from the Middle Pliocene to the late Pleistocene in Italy. The state of the art. Rivista Italiana di Paleontologia e Stratigrafia, 103, 369–388.Google Scholar
  41. Guillén Castejón, J. (2010). Canal Negre 1, un jaciment càrstic de vertebrats del Miocè, Pliocè i Pleistocè de Catalunya. Exploracions, 19, 7–87.Google Scholar
  42. Haas, G. (1951). The fauna of layer B of the Abu Usba Cave. In M. Stekelis & G. Haas (Eds.), The Abu Usba Cave (Mount Carmel). Israel Exploration Journal, 2, 35–47.Google Scholar
  43. Haas, G. (1967). Bemerkungen über die Fauna der Geula-Höhle, Carmel. Quaternaria, 9, 97–104.Google Scholar
  44. Heller, J. (1978). The faunal remains of Iraq e Zigan, a late Pleistocene site on Mt. Carmel. Israel Journal of Zoology, 27, 11–19.Google Scholar
  45. Hodrová, M. (1985). Amphibia of Pliocene and Pleistocene Včeláre localities (Slovakia). Casopis Mineralogii Geologii, 30(2), 145–161.Google Scholar
  46. Holdhaus, K. (1912). Über die Coelopteren- und Molluskenfauna des Monte Gargano (unter besonderer Berücksichtigung der Adriatisfrage). Denkschriften der Kaiserlichen Akademie der Wissenschaften, Mathematisch-Naturwissenschaftliche Klasse, 87, 432–465.Google Scholar
  47. Holman, J. A. (1992). The Boxgrove, England, Middle Pleistocene herpetofauna: Paleogeographic, evolutionary, stratigraphic, and paleoecological relationships. Historical Biology, 6, 263–279.CrossRefGoogle Scholar
  48. Holman, J. A. (1993). British Quaternary herpetofaunas: a history of adaptations to Pleistocene disruptions. Herpetological Journal, 3, 1–7.Google Scholar
  49. Holman, J. A. (1998). Pleistocene amphibians and reptiles in Britain and Europe. Oxford Monographs on geology and geophysics. New York and Oxford: Oxford University Press.Google Scholar
  50. Iosif, R., Papeş, M., Samoilă, C., & Cogălniceanu, D. (2014). Climate-induced shifts in the niche similarity of two related spadefoot toads (genus Pelobates). Organisms Diversity and Evolution, 14, 397–408.CrossRefGoogle Scholar
  51. Jesse, R., Pfenninger, M., Fratini, S., Scalici, M., Streit, B., & Schubart, C. D. (2009). Disjunct distribution of the Mediterranean freshwater crab Potamon fluviatile—natural expansion or human introduction? Biological Invasions, 11, 2209–2221.CrossRefGoogle Scholar
  52. Karaman, I., Hammouti, N., Pavićević, D., Kiefer, A., Horvatović, M., & Seitz, A. (2011). The genus Troglophilus Krauss, 1879 (Orthoptera: Rhaphidophoridae) in the west Balkans. Zoological Journal of the Linnean Society, 163, 1035–1063.CrossRefGoogle Scholar
  53. Kobelt, W. (1898). Studien zur Zoogeographie. II. Die Fauna der meridionalen Sub-Region. Wiesbaden: Kreidel.Google Scholar
  54. Korábek, O., Juřičková, L., & Petrusek, A. (2014). Resurrecting Helix straminea, a forgotten escargot with trans-Adriatic distribution: first insights into the genetic variation within the genus Helix (Gastropoda: Pulmonata). Zoological Journal of the Linnean Society, 171, 72–91.CrossRefGoogle Scholar
  55. Lanza, B., Andreone, F., Bologna, M. A., Corti, C., & Razzetti, E. (2007). Fauna d’Italia. Amphibia, XLII. Bologna: Calderini.Google Scholar
  56. López-García, J. M., Luzi, E., Berto, C., Peretto, C., & Arzarello, M. (2015). Chronological context of the first hominin occurrence in southern Europe: the Allophaiomys ruffoi (Arvicolinae, Rodentia, Mammalia) from Pirro 13 (Pirro Nord, Apulia, southwestern Italy). Quaternary Science Reviews, 107, 260–266.CrossRefGoogle Scholar
  57. Manzano, A. (2015). Les amphibiens et les reptiles des sites Pléistocène moyen et supérieur de la France méditerranéenne (Caune de l’Arago, grotte du Lazaret et Baume Moula-Guercy). Étude systématique, reconstitutions paléoclimatiques et paléoenvironnementales. Unpublished PhD thesis, Université de Perpignan Via Domitia, Perpignan.Google Scholar
  58. Martín, C., & Sanchiz, B. (2015). Lisanfos KMS. Version 1.2. Museo Nacional de Ciencias Naturales, MNCN-CSIC, Madrid, Spain, Online reference accessible at: Accessed 10 Feb 2015.
  59. Masini, F., & Sala, B. (2007). Large and small-mammal distribution patterns and chronostratigraphic boundaries from the Late Pliocene to the Middle Pleistocene of the Italian peninsula. Quaternary International, 160, 43–56.CrossRefGoogle Scholar
  60. Młynarski, M. (1977). New notes on the amphibian and reptilian fauna of the Polish Pliocene and Pleistocene. Acta Zoologica Cracoviensia, 22, 13–36.Google Scholar
  61. Napoleone, G., Albianelli, A., Azzaroli, A., Bertini, A., Magi, M., & Mazzini, M. (2003). Calibration of the upper Valdarno Basin to the Plio-Pleistocene for correlating the Apennine continental sequences. Il Quaternario, 16, 131–166.Google Scholar
  62. Paunovic, M., & Dimitrijevic, V. (1990). Gornjopleistocenska fauna nižih Vertebrata iz Smolćke pećine u jugozapadnoj Srbiji. Rad Jugoslavenske Akademije Znanosti Umjetnosti, 24(449), 77–87.Google Scholar
  63. Pavia, G., Bertok, C., Ciampo, G., Di Donato, V., Martire, L., Masini, F., Pavia, M., Santangelo, N., Taddei Ruggiero, E., & Zunino, M. (2010). Tectono-sedimentary evolution of the Pliocene to lower Pleistocene succession of the Apricena-Lesina-Poggio Imperiale quarrying district (western Gargano, southern Italy). Bollettino della Societa Geologica Italiana, 129, 132–155.Google Scholar
  64. Pavia, M., Zunino, M., Coltorti, M., Angelone, C., Arzarello, M., Bagnus, C., Belluci, L., Colombero, S., Marcolini, F., Peretto, C., Petronio, C., Petrucci, M., Pieruccini, P., Sardella, R., Tema, E., Villier, B., & Pavia, G. (2012). Stratigraphical and paleontological data from the early Pleistocene Pirro 10 site of Pirro Nord (Puglia, South eastern Italy). Quaternary International, 267, 40–55.CrossRefGoogle Scholar
  65. Petrucci, M., Cipullo, A., Martínez-Navarro, B., Rook, L., & Sardella, R. (2013). The late Villafranchian (Early Pleistocene) carnivores (Carnivora, Mammalia) from Pirro Nord (Italy). Palaeontographica Abteilung A, 298, 113–146.CrossRefGoogle Scholar
  66. Peyrouse, J. B., & Marquet, J. C. (2010). Les amphibiens et les reptiles du Pléistocène supérieur ancien de la Roche-Cotard III (Indre-et-Loire). Bulletin de la Société d’histoire naturelle d’Antun, 199, 37–44.Google Scholar
  67. Picarello, O., & Scillitani, G. (1988). Genetic distances between the populations of Cyrtodactylus kotschyi (Squamata: Gekkonidae) from Apulia and Greece. Amphibia–Reptilia, 9, 245–250.Google Scholar
  68. Ratnikov, V. Y. (2001). Pliocene anurans of East-European platform. Russian Journal of Herpetology, 8(3), 171–178.Google Scholar
  69. Ratnikov, V. Y. (2009). Fossil remains of modern amphibian and reptile species as the material for studing of their areas history. Science Research Works of the Geological Institute of Voronezh, 59, 1–91 (in Russian).Google Scholar
  70. Roček, Z. (1981). Cranial anatomy of frogs of the family Pelobatidae Stannius, 1856, with outlines of their phylogeny and systematics. Acta Universitatis Carolinae Biologica, 1980(1–2), 1–164.Google Scholar
  71. Roček, Z. (1988). List of fossil amphibians of Czechoslovakia. Acta Zoologica Cracoviensia, 31(19), 513–540.Google Scholar
  72. Roček, Z. (2013). Mesozoic and tertiary anura of Laurasia. In J.D. Gardner & R.L. Nydam (Eds), Mesozoic and Cenozoic lissamphibian and squamate assemblages of Laurasia. Palaeobiodiversity and Palaeoenvironments, 93, 397–439.Google Scholar
  73. Roček, Z., & Wuttke, M. (2010). Amphibia of Enspel (late Oligocene, Germany). Palaeobiodiversity and Palaeoenvironments, 90, 321–340.CrossRefGoogle Scholar
  74. Roček, Z., Eaton, J. E., Gardner, J. D., & Přikryl, T. (2010). Evolution of anuran assemblages in the Late Cretaceous of Utah, USA. Palaeobiodiversity and Palaeoenvironments, 90, 341–393.CrossRefGoogle Scholar
  75. Roček, Z., Wuttke, M., Gardner, J. D., & Singh Bhullar, B.-A. (2014). The Euro-American genus Eopelobates, and a re-definition of the family Pelobatidae (Amphibia, Anura). Palaeobiodiversity and Palaeoenvironments, 94, 529–567.CrossRefGoogle Scholar
  76. Rook, L., & Sardella, R. (2005). Hystrix refossa Gervais 1852, from Pirro Nord, early Pleistocene, southern Italy. Rivista Italiana di Paleontologia e Stratigrafia, 111, 485–492.Google Scholar
  77. Rook, L., & Sardella, R. (2013). New data on the early Pleistocene large sized porcupine from Pirro Nord (Apricena, Apulia, Italy). Palaeontographica Abteilung A, 298, 87–94.CrossRefGoogle Scholar
  78. Salari, L., Kotsakis, T., & Petronio, C. (2013). Early Pleistocene bats from Pirro Nord (Apulia, southern Italy). Palaeontographica Abteilung A, 298, 55–72.CrossRefGoogle Scholar
  79. Sanchiz, B. (1998). Salientia. In P. Wellnhofer (Ed.), Encyclopedia of paleoherpetology, part 4 (pp. 1–275). Munich: Verlag Dr. Friedrich Pfeil.Google Scholar
  80. Sanchíz, F., & Młynarski, M. (1979). Remarks on the fossil anurans from the Polish Neogene. Acta Zoologica Cracoviensia, 24, 153–174.Google Scholar
  81. Schmalfuss, H., Paragamian, K., & Sfenthourakis, S. (2004). The terrestrial isopods (Isopoda: Oniscidea) of Crete and the surrounding islands. Stuttgarter Beiträge zur Naturkunde. Serie A (Biologie), 662, 1–74.Google Scholar
  82. Scillitani, G., Turrisi, G. F., & Vaccaro, A. (2009). Zamenis situla (Linnaeus, 1758). In R. Sindaco, G. Doria, E. Razzetti, & F. Bernini (Eds.), Atlas of Italian amphibians and reptiles (pp. 584–587). Edizioni Polistampa. Firenze: Societas Herpetologica Italica.Google Scholar
  83. Shpun, S., Hoffman, J., Nevo, E., & Katz, U. (1993). Is the distribution of Pelobates syriacus related to its limited osmoregulatory capacity? Comparative Biochemistry and Physiology A: Physiology, 105(1), 135–139.CrossRefGoogle Scholar
  84. Sillero, N., Campos, J., Bonardi, A., Corti, C., Creemers, R., Crochet, P.-A., Isailovic, J. C., Denoël, M., Ficetola, G. F., Gonçalves, J., Kuzmin, S., Lymberakis, P., de Pous, P., Rodríguez, A., Sindaco, R., Speybroeck, J., Toxopeus, B., Vieites, D. R., & Vences, M. (2014). Updated distribution and biogeography of amphibians and reptiles of Europe. Amphibia–Reptilia, 35, 1–264.Google Scholar
  85. Sindaco, R., Doria, G., Razzetti, E., & Bernini, F. (2006). Atlante degli Anfibi e dei Rettili d’Italia. Atlas of Italian amphibians and reptiles. Edizioni Polistampa. Firenze: Societas Herpetologica Italica.Google Scholar
  86. Sinsch, U. (1991). Mini-review: the orientation behaviour of amphibians. Herpetological Journal, 1, 541–544.Google Scholar
  87. Stănescu, F., Iosif, R., Székely, D., Székely, P., Roşioru, D., & Cogălniceanu, D. (2013). Salinity tolerance in Pelobates fuscus (Laurenti, 1768) tadpoles (Amphibia: Pelobatidae). Travaux du Muséum National d’Histoire Naturelle “Gringore Antipa”, 56(1), 103–108.Google Scholar
  88. Tarkhnishvili, D., Serbinova, I., & Gavashelishvili, A. (2009). Modelling the range of Syrian spadefoot toad (Pelobates syriacus) with combination of GIS-based approaches. Amphibia-Reptilia, 30, 401–412.CrossRefGoogle Scholar
  89. Valla, F. R., Bar-Yosef, O., Smith, P., Desse, J., & Tchernov, E. (1986). Un nouveau sondage sur la terrace d’El Ouad, Israel (1980-1981). Paléorient, 12, 21–38.CrossRefGoogle Scholar
  90. van Andel, T. H., & Shackleton, J. C. (1982). Late paleolithic and mesolithic coastlines of Greece and the Aegean. Journal of Field Archeology, 9, 445–454.Google Scholar
  91. Velić, J., & Malvić, T. (2011). Depositional conditions during Pliocene and Pleistocene in Northern Adriatic and possible lithostratigraphic division of these rocks. Nafta, 62(1–2), 25–32.Google Scholar
  92. Venczel, M. (2004). Middle Miocene anurans from the Carpathian Basin. Palaeontographica Abteilung A, 271, 151–174.Google Scholar
  93. Venczel, M., & Hír, J. (2013). Amphibians and squamates from the Miocene of Felsötárkány Basin, N-Hungary. Palaeontographica Abteilung A, 300, 117–158.Google Scholar
  94. Vergnaud-Grazzini, C. (1970). Les amphibiens fossiles du gisement d’Arondelli. Palaeontolographia Italica, 66, 47–65.Google Scholar
  95. Wagler, J. (1830). Natürliches System der Amphibien, mit vorangehender Classification der Säugthiere und Vögel. Ein Beitrag zur vergleichenden Zoologie. München: J. G. Cotta.Google Scholar

Copyright information

© Senckenberg Gesellschaft für Naturforschung and Springer-Verlag Berlin Heidelberg 2015

Authors and Affiliations

  • Hugues-Alexandre Blain
    • 1
    • 2
  • Massimo Delfino
    • 3
    • 4
  • Claudio Berto
    • 5
    • 6
  • Marta Arzarello
    • 5
    • 6
  1. 1. Institut Català de Paleoecologia Humana i Evolució Social (IPHES)TarragonaSpain
  2. 2.Area de PrehistoriaUniversitat Rovira i Virgili (URV)TarragonaSpain
  3. 3.Dipartimento di Scienze della TerraUniversità di TorinoTorinoItaly
  4. 4.Institut Català de Paleontologia Miquel Crusafont (ICP)Universitat Autònoma de Barcelona (UAB)Cerdanyola del VallèsSpain
  5. 5.Dipartimento di Studi Umanistici, Sezione di Scienze Preistoriche e AntropologicheUniversità degli Studi di FerraraFerraraItaly
  6. 6.LT TekneHubFerraraItaly

Personalised recommendations