Monitor lizards (genus Varanus) inhabited Europe at least from the early Miocene to the Pleistocene. Their fossil record is limited to about 40 localities that have provided mostly isolated vertebrae. Due to the poor diagnostic value of these fossils, it was recently claimed that all the European species described prior to the 21st century are not taxonomically valid and a new species, Varanus amnhophilis, was erected on the basis of fragmentary material including cranial elements, from the late Miocene of Samos (Greece). We re-examined the type material of Varanus marathonensis Weithofer, 1888, based on material from the late Miocene of Pikermi (Greece), and concluded that it is a valid, diagnosable species. Previously unpublished Iberian material from the Aragonian (middle Miocene) of Abocador de Can Mata (Vallès-Penedès Basin, Barcelona) and the Vallesian (late Miocene) of Batallones (Madrid Basin) is clearly referable to the same species on a morphological basis, further enabling to provide an emended diagnosis for this species. Varanus amnhophilis appears to be a junior subjective synonym of V. marathonensis. On the basis of the most complete fossil Varanus skeleton ever described, it has been possible to further resolve the internal phylogeny of this genus by cladistically analyzing 80 taxa coded for 495 morphological and 5729 molecular characters. Varanus marathonensis was a large-sized species distributed at relatively low latitudes in both southwestern and southeastern Europe from at least MN7+8 to MN12. Our cladistic analysis nests V. marathonensis into an eastern clade of Varanus instead of the African clade comprising Varanus griseus, to which it had been related in the past. At least two different Varanus lineages were present in Europe during the Neogene, represented by Varanus mokrensis (early Miocene) and V. marathonensis (middle to late Miocene), respectively.

ARAID Fundación Conjunto Paleontológico de Teruel Dinópolis Teruel Spain

Departamento de Paleobiología Museo Nacional de Ciencias Naturales Consejo Superior de Investigaciones Cientificas Madrid Spain

Department of Geological Sciences Faculty of Science Masaryk University Brno Czech Republic

Dipartimento di Scienze della Terra Università di Torino Torino Italy

Division of Anthropology American Museum of Natural History New York United States of America

Institut Català de Paleontologia Miquel Crusafont Universitat Autònoma de Barcelona Cerdanyola del Vallès Barcelona Spain

Laboratory of Geology and Paleontology Department of Geology Aristotle University of Thessaloniki Thessaloniki Greece

New York Consortium in Evolutionary Primatology New York United States of America

School of Earth and Environmental Sciences University of Manchester Manchester United Kingdom

School of Earth Sciences University of Bristol Bristol United Kingdom

Universidad Estatal de la Peninsula de Santa Elena La Libertad Santa Elena Ecuador

See more in PubMed

Pianka ER, King DR, Allen King R. Varanoid Lizards of the World. Bloomington: Indiana University Press; 2004.

Holmes RB, Murray AM, Attia YS, Simons EL, Chatrath P. Oldest known Varanus (Squamata: Varanidae) from the Upper Eocene and Lower Oligocene of Egypt: support for an African origin of the genus. Palaeontology. 2010; 53: 1099–1110.

Rage JC, Bailón S. Amphibians and squamate reptiles from the late early Miocene (MN 4) of Béon 1 (Montréal-du-Gers), southwestern France. Geodiversitas 2005; 27: 413–441.

Georgalis GL, Villa A, Delfino M. The last European varanid: demise and extinction of monitor lizards (Squamata, Varanidae) from Europe. J Vertebr Paleontol. 2017; 37: e1301946.

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

Augé ML, Guével B. New varanid remains from the Miocene (MN4–MN5) of France: inferring fossil lizard phylogeny from subsets of large morphological data sets. J Vertebr Paleontol. 2018; 38: e1410483.

Estes R. Sauria terrestria, Amphisbaenia. Stuttgart: G Fischer; 1983.

Böhme M, Ilg A fosFARbase; 2003 [accessed 2012 March 1]. Available from: http://www.wahre-staerke.com/

Molnar RE. The long and honourable history of monitors and their kin In: Pianka ER, King DR, Allen King R, editors. Varanoid Lizards of the World. Bloomington: Indiana University Press; 2004. pp. 10–67.

Conrad JL, Ast JC, Montanari S, Norell MA. A combined evidence phylogenetic analysis of Anguimorpha (Reptilia: Squamata). Cladistics. 2011; 27: 230–277. PubMed

Conrad JL, Balcarcel A, Mehling C. Earliest example of a giant monitor lizard (Varanus, Varanidae, Squamata). PLoS ONE. 2012; 7: e41767 10.1371/journal.pone.0041767 PubMed DOI PMC

Conrad JL, Balcarcel A, Mehling C. Miocene Asian invasion of Europe by Varanus (Varanidae). J Vertebr Paleontol. 2009; 29(Suppl): 81A.

Koufos GD, Kostopoulos DS, Vlachou TD. The Late Miocene mammal faunas of the Mytilinii Basin, Samos Island, Greece: new collection– 16. Biochronology. Beitr Paläontol. 2009; 31: 397–408.

Koufos GD. The Neogene mammal localities of Greece: faunas, chronology and biostratigraphy. Hell J Geosci. 2006; 41: 183–214.

Weithofer A. Beiträge zur Kenntniss der Fauna von Pikermi bei Athen. Beitr Paläontol Geol Österr-Ung Orients. 1888; 6: 225–292.

Delfino M, Alba DM, Carmona R, Luján AH, Robles JM. European monitor lizards (Anguimorpha, Varanidae, Varanus): new materials and new perspectives. J Vertebr Paleontol. 2011; 31(Suppl), pp. 97–98.

Delfino M, Alba DM, Luján ÀH, Carmona R, Abella J, Robles JM, et al. Varanus marathonensis Weithofer, 1888: morphology, systematics, and paleobiogeography of the European monitor lizards In: Devleeschouwer X, Heyvaert VMA, Louwye S, Piessens K, Smith T, editors. Moving Plates and Melting Icecaps–Processes and Forcing Factors in Geology. Brussels: Royal Belgian Institute of Natural Sciences; 2012. p. 156.

Goloboff PA, Catalano SA. TNT version 1.5, including a full implementation of phylogenetic morphometrics. Cladistics. 2016; 32: 221–238. PubMed

Brochu CA. Alligatorine phylogeny and the status of Allognatosuchus Mook, 1921. J Vertebr Paleontol. 2004; 24: 857–873.

Alba DM, Casanovas-Vilar I, Garcés M, Robles JM. Ten years in the dump: an updated review of the Miocene primate-bearing localities from Abocador de Can Mata (NE Iberian Peninsula). J Hum Evol. 2017; 102: 12–20. 10.1016/j.jhevol.2016.09.012 PubMed DOI

Cabrera L, Calvet F, Guimerà J, Permanyer A. El registro sedimentario miocénico en los semigrábens del Vallès-Penedès y de El Camp: organización secuencial y relaciones tectónica sedimentación. In: Colombo F, editor. I Congreso del Grupo Español del Terciario, Vic 1991. Libro-Guía Excursion n° 4. Barcelona: Universitat de Barcelona. 132 pp.

Cabrera L, Roca E, Garcés M, Porta J de. Estratigrafía y evolución tectonosedimentaria oligocena superior-neógena del sector central del margen catalán (Cadena Costero-Catalana) In: Vera JA, editor. Geología de España. Madrid: Instituto Geológico y Minero de España; 2004. pp. 569–573.

Cabrera L, Calvet F. Onshore Neogene record in NE Spain: Vallès-Penedès and El Camp half-grabens (NW Mediterranean) In: Friend P, Dabrio CJ, editors. Tertiary basins of Spain: the stratigraphic record of crustal kinematics. Cambridge: Cambridge University Press; 1996. pp. 97–105.

Roca E, Sans M, Cabrera L, Marzo M. Oligocene to Middle Miocene evolution of the central Catalan margin (northwestern Mediterranean). Tectonophysics. 1999; 315: 209–229.

Gibert JM de, Casanovas-Vilar I. Contexto geológico del Mioceno de la cuenca del Vallès-Penedès. Paleontol Evol, Mem espec. 2011; 6: 39–45.

Casanovas-Vilar I, Alba DM, Robles JM, Moyà-Solà S. Registro paleontológico continental del Mioceno de la cuenca del Vallès-Penedès. Paleontol Evol, Mem espec. 2011; 6: 55–80.

Casanovas-Vilar I, Madern A, Alba DM, Cabrera L, García-Paredes I, Van den Hoek Ostende LW, DeMiguel D, Robles JM, Furió M, Van Dam J, Garcés M, Angelone C, Moyà-Solà S. The Miocene mammal record of the Vallès-Penedès Basin (Catalonia). C R Palevol. 2016; 15: 791–812.

Alba DM, Moyà-Solà S, Casanovas-Vilar I, Galindo J, Robles JM, Rotgers C, et al. Los vertebrados fósiles del Abocador de Can Mata (els Hostalets de Pierola, l’Anoia, Cataluña), una sucesión de localidades del Aragoniense superior (MN6 y MN7+8) de la cuenca del Vallès-Penedès–Campañas 2002–2003, 2004 y 2005. Estud Geol. 2006; 62: 295–312.

Alba DM, Robles JM, Rotgers C, Casanovas-Vilar I, Galindo J, Moyà-Solà S, et al. Middle Miocene vertebrate localities from Abocador de Can Mata (els Hostalets de Pierola, Vallès-Penedès Basin, Catalonia, Spain): an update after the 2006–2008 field campaigns. Paleolusitana. 2009; 1: 59–73.

Alba DM, Casanovas-Vilar I, Robles JM, Moyà-Solà S. Parada 3 –El Aragoniense superior y la transición con el Vallesiense: Can Mata y la exposición paleontologica de els Hostalets de Pierola. Paleontol Evol, Mem espec. 2011; 6: 95–109.

Agustí J, Cabrera L, Moyà-Solà S. Sinopsis estratigráfica del Neógeno de la fosa del Vallés-Penedés. Paleontol Evol. 1985; 18: 57–82.

Moyà-Solà S, Köhler M, Alba DM, Casanovas-Vilar I, Galindo J, Robles JM, et al. First partial face and upper dentition of the Middle Miocene hominoid Dryopithecus fontani from Abocador de Can Mata (Vallés-Penedés Basin, Catalonia, NE Spain): taxonomic and phylogenetic implications. Am J Phys Anthropol 2009; 139: 126–145. 10.1002/ajpa.20891 PubMed DOI

Crusafont M, Truyols J. Catálogo paleomastológico del Mioceno del Vallés-Penedés y de Calatayud-Teruel. Sabadell: Museo de la Ciudad de Sabadell; 1954.

Golpe-Posse JM. Faunas de yacimientos con suiformes en el Terciario español. Paleontol Evol. 1974; 8: 1–87.

Agustí J, Cabrera L, Garcés M, Parés JM. The Vallesian mammal succession in the Vallès-Penedès basin (northeast Spain): paleomagnetic calibration and correlation with global events. Paleogeogr Paleoclimatol Paleoecol. 1997; 133: 149–180.

Casanovas-Vilar I, Garcés M, Van Dam J, García-Paredes I, Robles JM, Alba DM. An updated biostratigraphy for the late Aragonian and the Vallesian of the Vallès-Penedès Basin (Catalonia). Geol Acta. 2016; 14: 195–217.

Casanovas-Vilar I, Alba DM, Garcés M, Robles JM, Moyà-Solà S. Updated chronology for the Miocene hominoid radiation in Western Eurasia. Proc Natl Acad Sci U S A. 2011; 108: 5554–5559. 10.1073/pnas.1018562108 PubMed DOI PMC

Morales J, Capitán J, Calvo JP, Sesé C. Nuevo yacimiento de vertebrados del Mioceno Superior al Sur de Madrid (Cerro Batallones, Torrejón de Velasco). Geogaceta. 1992; 12: 77–80.

Morales J, Alcalá L, Álvarez Sierra MA, Antón M, Azanza B, Calvo JP et al. Paleontología del sistema de yacimientos de mamíferos miocenos del Cerro de los Batallones, Cuenca de Madrid. Geogaceta. 2004; 35: 139–142.

Morales J, Pozo M, Silva PG, Domingo MS, López-Antoñanzas R, Alvarez Sierra MA et al. El sistema de yacimientos de mamíferos miocenos del Cerro de los Batallones, Cuenca de Madrid: estado actual y perspectivas In: Esteve J, Meléndez G, editors. Palaeontologica Nova. 2008. pp. 71–114.

Pozo M, Calvo JP, Silva PG, Morales J, Peláez-Campomanes P, Nieto M. Geología del sistema de yacimientos de mamíferos miocenos del Cerro de los Batallones, Cuenca de Madrid. Geogaceta. 2004; 35: 143–146.

Fraile S, Pesquero MD, García Somoza P, Morales J. El Cerro de los Batallones: un caso práctico de actuación paleontológica de carácter preventivo. Col Paleontol, Vol extr. 2003; 1: 163–169.

Calvo JP, Pozo M, Silva PG, Morales J. Pattern of sedimentary infilling of fossil mammal traps formed in pseudokarst at Cerro de los Batallones, Madrid Basin, central Spain. Sedimentology. 2013; 60: 1681–1708.

Domingo MS, Domingo L, Sánchez IM, Alberdi MT, Azanza B, Morales J. New insights on the taphonomy of the exceptional mammalian fossil sites of Cerro del los Batallones (Late Miocene, Spain) based on rare earth element geochemistry. Palaios. 2011; 26: 55–65.

López-Antoñanzas R, Peláez-Campomanes P, Álvarez-Sierra MA, García-Paredes I. New species of Hispanomys (Rodentia, Cricetodontinae) from the Upper Miocene of Batallones (Madrid, Spain). Zool J Linn Soc. 2010; 160: 725–747.

The NOW Community. New and Old Worlds Database of Fossil Mammals (NOW); 2018 [accessed 2018 July 21]. Available from: http://www.helsinki.fi/science/now/.

Koufos GD. Neogene mammal biostratigraphy and chronology of Greece In: Wang X, Flynn LJ, Fortelius M, editors, Fossil Mammals of Asia: Neogene Biostratigraphy and Chronology. New York: Columbia University Press; 2013. pp. 595–621.

Böhme M, Spassov N, Ebner M, Geraads D, Hristova L, Kirscher U et al. Messinian age and savannah environment of the possible hominin Graecopithecus from Europe. PLoS ONE. 2017; 12: e0177347 10.1371/journal.pone.0177347 PubMed DOI PMC

Gaudry A. Animaux fossiles et Géologie de l'Attique d'après les Recherches faites en 1855–56 et en 1860. Paris: F. Savy; 1862.

Fejérváry GJ de. Contributions to a monograph on fossil Varanidae and on Megalanidae. Ann Mus Natl Hung. 1918; 16: 341–467.

Bhullar BAS. Cranial osteology of Exostinus serratus (Squamata: Anguimorpha), fossil sister taxon to the enigmatic clade Xenosaurus. Zool J Linn Soc. 2010; 159: 921–953.

Hoffstetter R. Sur la position systématique de Necrosaurus: Saurien de l'Eocène européen. CR Soc Geol France 1954; 16: 422–424.

Maxwell EE, Caldwell MW, Lamoureux DO. The structure and phylogenetic distribution of amniote plicidentine. J. Vertebr Paleontol 2011; 31: 553–561.

Jollie MT. The head skeleton of the lizard. Acta Zool. 1960; 41: 1–64.

Rage JC, Augé M. Squamate reptiles from the middle Eocene of Lissieu (France). A landmark in the middle Eocene of Europe. Geobios 2010; 43: 253–268.

Bell CJ, Mead JI, Swift SL. Cranial osteology of Moloch horridus (Reptilia: Squamata: Agamidae). Rec West Aust Mus. 2009; 25: 201–237.

Georgalis GL, Rage JC, Bonis L de, Koufos GD. Lizards and snakes from the late Miocene hominoid locality of Ravin de la Pluie (Axios Valley, Greece). Swiss J Geosci. 2018; 111: 169–181.

Ast JC. Mitochondrial DNA evidence and evolution in Varanoidea (Squamata). Cladistics. 2001; 17: 211–226. PubMed

Gaudry A. Note sur les débris d'oiseaux et de reptiles trouvés à Pikermi, Grèce, suivie de quelques remarques de paléontologie générale. Bull Soc Geol Fr. 1862. b; 19: 629–640.

Gaudry A. Animaux fossiles et Géologie de l'Attique d'après les recherches faites en 1855–56 et en 1860 –Atlas. Paris: F Savy; 1862–1867.

Mertens R. Die Familie der Warane (Varanidae)–Zweiter Teil: der Schädel. Abh Senckenb Naturforsch Ges 1942; 465: 117–234.

Mertens R. Der Schädel von Varanus salvadorii In: Herre W, editor. Neue Ergebnisse und Probleme der Zoologie. Leipzig: Geest & Portig; 1950. pp.: 561–566.

Mertens R. Der Schädel des indischen Gelbwarans, Varanus flavescens. Senckenb Biol. 1959; 40: 113–115.

Conrad JL, Rieppel O, Grande L. Re-assessment of varanid evolution based on new data from Saniwa ensidens Leidy, 1870 (Squamata, Reptilia). Am Mus Novit. 2008; 3630: 1–15.

Conrad JL Phylogeny and systematics of Squamata (Reptilia) based on morphology. Bull Am Mus Nat Hist. 2008; 310: 1–182.

Clos LM. A new species of Varanus (Reptilia: Sauria) from the Miocene of Kenya. J Vertebr Paleontol. 1995; 15: 254–267.

Nopcsa F. Zur Kenntnis der fossilen Eidechsen. Beitr Paläontol Geol Österr-Ung Orients. 1908; 21: 33–62.

Hoffstetter R. Varanidae et Necrosauridae fossiles. Bull Mus natl Hist nat. 1943; 15: 134–141.

Rage JC, Sen S. Les amphibiens et les reptiles du Pliocène supérieur de Calta (Turquie). Géol Médit. 1976; 3: 127–134.

Bolkay J. Additions to the fossil herpetology of Hungary from the Pannonian and Praeglacial periode. Mitt Jahrb K Ung Geol Reichsanst. 1913; 21: 217–230.

Młynarski M. Lizard from the Pliocene of Poland. Acta Palaeontol Pol. 1956; 1: 135–152.

Roger O. Wirbelthierreste aus dem Dinotheriensande–III Theil. Ber naturwiss Ver Schwaben Neuburg. 1900; 34: 53–70.

Hoffstetter R. Presence de Varanidae (Reptilia, Sauria) dans le Miocene de Catalogne. Considerations sur l’histoire de la famille. Bull Mus natl Hist nat. 1969; 40: 1051–1064.

Alférez Delgado F, Brea López P. Estudio preliminar de los restos de peces, anfibios y reptiles del yacimiento mioceno de Córcoles (Guadalajara). Bol Real Soc Esp Hist Nat, Secc geol. 1981; 79: 5–20.

Tempfer PM. The herpetofauna (Amphibia: Caudata, Anura; Reptilia: Scleroglossa) of the Upper Miocene locality Kohfidisch (Burgenland, Austria). Beitr Paläontol. 2005; 29: 145–253.

Venczel M. Lizards from the Late Miocene of Polgárdi (W-Hungary). Nymphaea. 2006; 33: 25–38.

Zerova GA, Chkhikvadze VM. Neogene varanids of the URSS In: Roček Z, editor. Studies in Herpetology. Prague: Charles University; 1986. pp. 689–694.

Delfino M, Rage JC, Bolet A, Alba DM. Early Miocene dispersal of the lizard Varanus into Europe: reassessment of vertebral material from Spain. Acta Palaeontol Pol. 2013; 58: 731–735.

Morelli N. Resti organici rinvenuti nella caverna delle Arene Candide presso Finalmarina. Atti Soc ligust Sci nat geogr. 1891; 2: 171–205.

Delfino M. Erpetofaune italiane del Neogene e del Quaternario. Ph.D. dissertation. Modena: Università degli Studi di Modena e Reggio Emilia; 2002.

Amer SAM, Kumazawa Y. Timing of a mtDNA gene rearrangement and intercontinental dispersal of varanid lizards. Genes Genet Syst. 2008; 83: 275–280. PubMed

Head JJ, Barrett PM, Rayfield EJ. Neurocranial osteology and systematic relationships of Varanus (Megalania) prisca Owen, 1859 (Squamata: Varanidae). Zool J Linn Soc. 2009; 155: 445–457.

Arida E, Böhme W. The origin of Varanus: when fossils, morphology, and molecules alone are never enough. Biawak. 2010; 4: 117–124.

Stanner M. Varanus griseus In: Pianka ER, King DR, Allen King R, editors. Varanoid Lizards of the World. Bloomington: Indiana University Press; 2004. pp. 104–132.

Sindaco R, Jeremčenko V. The reptiles of the western Palearctic. Vol. 1 Latina: Belvedere; 2008.

Levshakova IU. [A new monitor lizard from the Lower Pliocene of Tadzhikistan]. Tr Zool Inst Akad Nauk SSSR. 1986; 157: 101–106.

Molnar RE, Pianka ER (2004) Biogeography and phylogeny of varanoids In: Pianka ER, King DR, Allen King R, editors. Varanoid Lizards of the World. Bloomington: Indiana University Press; pp. 68–76.

Fuller S, Baverstock P, King D. Biogeographic origins of goannas (Varanidae): a molecular perspective. Mol Phylogenet Evol. 1998; 9: 294–307. 10.1006/mpev.1997.0476 PubMed DOI

Smith KT, Bhullar BAS, Holroyd PA. Earliest African record of the Varanus stem-clade (Squamata: Varanidae) from the Early Oligocene of Egypt. J Vertebr Paleontol. 2008; 28: 909913.

Augé M. Evolution des lézards du Paléogène en Europe. Mém Mus Natl Hist Nat. 2005; 192: 1–369.

Estes R, Hutchison JH. Eocene lower vertebrates from Ellesmere Island, Canadian Arctic Archipelago. Palaeogeogr Palaeoclimatol Palaeoecol. 1980; 30: 325–347.

Georgalis GL, Villa A, Delfino M. First description of a fossil chamaeleonid from Greece and its relevance for the European biogeographic history of the group. Sci Nat. 2016; 103: 12. PubMed

Rage JC (1997) Paleobiological and paleogeographical background of the European herpetofauna In: Gasc JP, Cabela A, Crnobrnja-Isailovic J, Dolmen D, Grossenbacher K, et al., editors. Atlas of Amphibians and Reptiles in Europe. Paris: Muséum national d'Histoire naturelle; 1997. pp. 23–28.

Szyndlar Z, Rage JC. West Paleartic cobras of the genus Naja (Serpentes: Elapidae): interrelationships among extinct and extant species. Amphibia-Reptilia. 1990; 11: 385–400.

Delfino M, Rage JC, Rook L. Tertiary mammal turnover phenomena: What happened to the herpetofauna? Deinsea. 2003; 10: 153–161.

Rage JC, Roček Z. Evolution of anuran assemblages in the Tertiary and Quaternary of Europe, in the context of palaeoclimate and palaeogeography. Amphibia-Reptilia 2003; 24: 133–177.

Szyndlar Z, Schleich HH. Description of Miocene snakes from Petersbuch 2 with comments on the lower and middle Miocene ophidian faunas of southern Germany. Stutt Beitr Naturkd Ser B. 1993; 192: 1–47.

Ivanov M. Snakes of the lower/middle Miocene transition at Vieux-Collonges (Rhône, France), with comments on the colonisation of western Europe by colubroids. Geodiversitas. 2000; 22: 559–588.

Oaks JR. A time-calibrated species tree of Crocodylia reveals a recent radiation of the true crocodiles. Evolution 2011; 65: 3285–3297. 10.1111/j.1558-5646.2011.01373.x PubMed DOI

Delfino M, Böhme M, Rook L. First European evidence for transcontinental dispersal of Crocodylus (late Neogene of southern Italy). Zool J Linn Soc. 2007; 149: 293–307.

Rögl F. Circum-Mediterranean Miocene paleogeography In: Rössner GE, Heissig K, editors. The Miocene land mammals of Europe. Munich: F Pfeil; 1999. pp. 39–48.

Sen S. Dispersal of African mammals in Eurasia during the Cenozoic: ways and whys. Geobios. 2013; 46: 159–172.

López-Antoñanzas R. Neogene Ctenodactylidae, Thryonomyidae, and Zapodidae (Rodentia) from the Middle East: systematics, phylogeny, biostratigraphy, palaeogeography, and palaeoecology. Ph.D. dissertation. Paris: Muséum national d'Histoire naturelle; 2004

Harrison T. Miocene primates In: Fuentes A, editor. The International Encyclopedia of Primatology. Vol. 2 Chichester: J Wiley & Sons; 2017; pp. 809–813.

Böhme M. The Miocene climatic optimum: evidence from ectothermic vertebrates of Central Europe. Paleogeogr Paleoclimatol Paleoecol. 2003; 195: 389–401.

Bailón S, Blain HA. Faunes de reptiles et changements climatiques en Europe occidentale autour de la limite Plio-Pléistocène. Quaternaire. 2007; 18: 55–63.

Bailón S. Escamosos (Reptilia) fósiles del yacimiento de Moreda (Plioceno medio/superior, Granada, España). Rev Esp Paleontol, Num Extra. 1992; 3: 11–15.

Delfino M. Blanus from the Early Pleistocene of Southern Italy: another small tessera from a big mosaic In: Böhme W, Bischoff W, Ziegler T, editors. Herpetologia Bonnensis. Bonn: Societas Europaea Herpetologica; 1997. pp. 89–97.

Delfino M. A Pleistocene amphisbaenian from Sicily. Amphibia-Reptilia. 2003; 24: 407–414.

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

Colombero S, Angelone C, Bonelli E, Carnevale G, Cavallo O, Delfino M, et al. The upper Messinian assemblages of fossil vertebrate remains of Verduno (NW Italy): another brick for a latest Miocene bridge across the Mediterranean. N Jahrb Geol Paläontol Abh. 2014; 272: 287–324.

Colombero S, Alba DM, D'Amico C, Delfino M, Esu D, Giuntelli P, et al. Late Messinian mollusks and vertebrates from Moncucco Torinese, north-western Italy. Paleoecological and paleoclimatological implications. Palaeontol Electron. 2017; 20: PE 20.1.10.

Blain HA, Bailón S, Agustí J. The geographical and chronological pattern of herpetofaunal Pleistocene extinctions on the Iberian Peninsula. C R Palevol. 2016; 15: 731–744.