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Early Pliocene Varanus (Squamata, Varanidae) remains from Megalo Emvolon, Thessaloniki, Greece.

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Τίτλος: Early Pliocene Varanus (Squamata, Varanidae) remains from Megalo Emvolon, Thessaloniki, Greece.
Συγγραφείς: Drakopoulou C; Laboratory of Geology and Palaeontology, School of Geology, Aristotle University of Thessaloniki, Thessaloniki, Greece., Georgalis GL; Institute of Systematics and Evolution of Animals, Polish Academy of Sciences, Kraków, Poland., Lazaridis G; Laboratory of Geology and Palaeontology, School of Geology, Aristotle University of Thessaloniki, Thessaloniki, Greece., Kostopoulos DS; Laboratory of Geology and Palaeontology, School of Geology, Aristotle University of Thessaloniki, Thessaloniki, Greece.
Πηγή: Anatomical record (Hoboken, N.J. : 2007) [Anat Rec (Hoboken)] 2026 Aug; Vol. 309 (8), pp. 2100-2123. Date of Electronic Publication: 2025 Oct 15.
Τύπος έκδοσης: Journal Article
Γλώσσα: English
Στοιχεία περιοδικού: Publisher: John Wiley & Sons Country of Publication: United States NLM ID: 101292775 Publication Model: Print-Electronic Cited Medium: Internet ISSN: 1932-8494 (Electronic) Linking ISSN: 19328486 NLM ISO Abbreviation: Anat Rec (Hoboken) Subsets: MEDLINE
Imprint Name(s): Original Publication: Hoboken, NJ : John Wiley & Sons, 2007-
Ιατρικοί όροι (MeSH): Lizards*/anatomy & histology , Lizards*/classification , Skull*/anatomy & histology , Fossils*, Tooth/anatomy & histology ; Animals ; Greece ; Biological Evolution
Περίληψη: This study describes new fossil varanid material from a recently discovered fossil spot (MVL site) at the Megalo Emvolon locality near Thessaloniki, Northern Greece, dated to 4.2-3.2 Ma (Early Pliocene, late Ruscinian, MN 15). The fossils, likely representing a single individual, are referred to Varanus cf. marathonensis based on cranial features, particularly the parietal, frontal, postorbitofrontal, dentary, and teeth. A key diagnostic trait is the wide angle between the squamosal and parietal processes of the postorbitofrontal, distinguishing it from Varanus mokrensis from the Early Miocene of Central Europe. Comparative analysis shows closer morphological resemblance with extant Australasian taxa, especially Varanus flavescens, rather than African ones, suggesting an origin from an Asian lineage. If indeed conspecific with Varanus marathonensis, the occurrence of Megalo Emvolon could represent the latest confirmed record of the species in Europe highlighting its resilience and extending its known temporal range. Additionally, the fossils provide palaeoclimatic evidence, indicating a warm and humid environment during the Early Pliocene in northern Greece. Trends of size reduction in the Plio-Pleistocene Greek Varanus likely correlate with climatic changes following the end of the Miocene.
(© 2025 The Author(s). The Anatomical Record published by Wiley Periodicals LLC on behalf of American Association for Anatomy.)
References: Andreetto, F., Matsubara, K., Beets, C. J., Fortuin, A. R., Flecker, R., & Krijgsman, W. (2021). High Mediterranean water‐level during the Lago‐Mare phase of the Messinian salinity crisis: Insights from the Sr isotope records of Spanish marginal basins (SE Spain). Palaeogeography, Palaeoclimatology, Palaeoecology, 562, 110139. https://doi.org/10.1016/j.palaeo.2020.110139.
Arambourg, C., & Piveteau, J. (1929). Les Vertébrés du Pontien de Salonique. Annales de Paleontologie, 18, 1–139.
Arida, E., & Böhme, W. (2010). The origin of Varanus: when fossils, morphology, and molecules alone are never enough. Biawak, 4, 117–124.
Augé, M. L. (2005). Évolution des lézards du Paléogène en Europe (p. 369). Muséum national d'Histoire naturelle.
Augé, M. L., Folie, A., Smith, R., Phélizon, A., Gigase, P., & Smith, T. (2022). Revision of the oldest varanid, Saniwa orsmaelensis Dollo, 1923, from the earliest Eocene of northwest Europe. Comptes Rendus Palevol, 21, 511–529. https://doi.org/10.5852/cr-palevol2022v21a25.
Bailon, S. (1991). Amphibiens et Reptiles du Pliocène et du Quaternaire de France et d'Espagne: mise en place et évolution des faunes. Ph.D. dissertation, Paris: Université de Paris VII.
Bailon, S. (1992). Escamosos (Reptilia) fósiles del yacimiento de Moreda (Plioceno medio/superior, Granada, España). Revista Española de Paleontología, 7(3), 11–15.
Bailon, S., & Blain, H.‐A. (2007). Faunes de reptiles et changements climatiques en Europe Occidentale autour de la limite Plio‐Pléistocène. Quaternaire, 18(1), 55–63.
Berkovitz, B., & Shellis, P. (2017). Reptiles 1: Tuatara and lizards. In B. Berkovitz & P. Shellis (Eds.), The teeth of non‐mammalian vertebrates (pp. 153–200). Academic Press. https://doi.org/10.1016/C2014-0-02210-1.
Bochaton, C., Ivanov, M., & Claude, J. (2019). Osteological criteria for the specific identification of monitor lizards (Varanus Merrem, 1820) remains in subfossil deposits of Sundaland and continental Southeast Asia. Amphibia‐Reptilia, 40(2), 219–232. https://doi.org/10.1163/15685381-20181101.
Böhme, W., Ahmed, S. H., Al‐Sheikhly, O. F., Ararat, K., Auer, M., Khudur, F. A., & Langner, C. (2023). Desert monitor lizards (Squamata: Varanidae: Varanus: Psammosaurus) from the Middle East: further records of Nesterov's desert monitor, Varanus (P.) nesterovi Böhme, Ehrlich, Milto, Orlov et Scholz, 2015, from Iraq, and adjacent localities of Varanus (P.) g. griseus (Daudin, 1803) and Varanus (P.) g. caspius (Eichwald, 1831), with comments on biogeography and taxonomy. Russian Journal of Herpetology, 30, 518–528.
Bolkay, J. (1913). Additions to the fossil herpetology of Hungary from the Pannonian and Praeglacial periode. Mitteilungen Aus Dem Jahrbuche der königlich Ungarischen Geologischen Reichsanstalt, 21(7), 217–230.
Bosc, L. A. G. 1792). Observations sur l'Acheta sylvestris et Locusta punctatissima. Actes de la Société d'Histoire Naturelle de Paris, 1, 44.
Boulenger, G. A. (1885). Catalogue of the lizards in the British museum (natural history) (Vol. 2, p. 497). British Museum.
Boulenger, G. A. (1898). Third report on additions to the lizard collection in the natural‐history Museum1. Proceedings of the Zoological Society of London, 66(4), 912–923. https://doi.org/10.1111/j.1096-3642.1898.tb03194.x.
Čerňanský, A., & Patnaik, Ρ. (2026). The first cranial material of Varanus from the Miocene of India. The Anatomical Record, 309(3), 634–641. https://doi.org/10.1002/ar.25676.
Čerňanský, A., Smith, R., Smith, T., & Folie, A. (2024). Timing of intercontinental faunal migrations: Anguimorph lizards from the earliest Eocene (MP 7) of Dormaal, Belgium. Zoological Journal of the Linnean Society, 201, zlae082.
Čerňanský, A., Syromyatnikova, E. V., & Jablonski, D. (2018). The first record of amphisbaenian and anguimorph lizards (Reptilia, Squamata) from the upper Miocene Solnechnodolsk locality in Russia. Historical Biology, 32(7), 869–879. https://doi.org/10.1080/08912963.2018.1539973.
Čerňanský, A., Tabuce, R., & Vidalenc, D. (2025). The first evidence of iguanians and scincoids from the lower Eocene of the cos locality (Phosphorites du Quercy, France). Journal of Paleontology, 99, 1–440. https://doi.org/10.1017/jpa.2025.10109.
Clos, L. M. (1995). A new species of Varanus (Reptilia: Sauria) from the Miocene of Kenya. Journal of Vertebrate Paleontology, 15, 254–267.
Conrad, J. L., Balcarcel, A. M., & Mehling, C. M. (2012). Earliest example of a giant monitor lizard (Varanus, Varanidae, Squamata). PLoS One, 7(8), 1–14. https://doi.org/10.1371/journal.pone.0041767.
Crespo, E. G. (2001). Paleo‐herpetofauna de Portugal. Museu Bocage.
Daudin, F. M. (1802). Histoire Naturelle, génerale et particulière des reptiles, ouvrage faisant suite, à l'Histoire Naturelle générale et particulière, composée par Leclerc de Buffon, et rédigée par C.S. Sonnini. Tome troisième (p. 452). F. Dufart.
Daudin, F. M. (1803). Histoire naturelle génerale et particulière des reptiles; Ouvrage faisant suite aux Cuvres de Leclerc de Buffon, et partie du cours complet d'histoire naturelle rédigé par C.S. Sonnini, membre de plusieurs sociétés savantes. Tome huitième (p. 439). F. Dufart. pls. 93–100.
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(4), 731–735.
Dong, L., Wang, Y.‐Q., Zhao, Q., Vasilyan, D., Wang, Y., & Evans, S. E. (2022). A new stem‐varanid lizard (Reptilia, Squamata) from the early Eocene of China. Philosophical Transactions of the Royal Society B, 377, 20210041.
Estes, R. (1983). Handbuch der Paläoherpetologie 10A. Friedrich Pfeil, München.
Estes, R., de Queiroz, K., & Gauthier, J. A. (1988). Phylogenetic relationships within Squamata. In R. Estes & G. K. Pregill (Eds.), Phylogenetic relationships of the lizard families: Essays commemorating C.L. camp (pp. 119–281). Stanford University Press.
Fejérváry, G. J. (1918). Contributions to a monography on fossil Varanidae and on Megalanidae. Annales Historico‐Naturales Musei Nationalis Hungarici, 16, 341–467.
Fürbringer, M. (1900). Zur vergleichenden anatomie des Brustschulterapparates und der Schultermuskeln. Jenaische Zeitschrift für Naturwissenschaft, 34, 15–718.
Georgalis, G. L. (2019). Poor but classic: The squamate fauna from the late Miocene of Pikermi, near Athens, Greece. Comptes Rendus Palevol, 18(7), 801–815. https://doi.org/10.1016/j.crpv.2019.09.004.
Georgalis, G. L., Abdel Gawad, M. K., Hassan, S. M., El‐Barkooky, A. N., & Hamdan, M. A. (2020). Oldest co‐occurrence of Varanus and Python from Africa–first record of squamates from the early Miocene of Moghra formation, Western Desert, Egypt. PeerJ, 8, e9092. https://doi.org/10.7717/peerj.9092.
Georgalis, G. L., Čerňanský, A., & Klembara, J. (2021). Osteological atlas of new lizards from the Phosphorites du Quercy (France), based on historical, forgotten, fossil material. Geodiversitas, 43(9), 219–293. https://doi.org/10.5252/geodiversitas2021v43a9.
Georgalis, G. L., & Delfino, M. (2022a). The fossil record of lizards and snakes (Reptilia: Squamata) in Greece. In E. Vlachos (Ed.), Fossil vertebrates of Greece Vol. 1 ‐ basal vertebrates, amphibians, reptiles, Afrotherians, Glires, and primates (pp. 205–235). Springer Nature. https://doi.org/10.1007/978-3-030-68398-6_7.
Georgalis, G. L., & Delfino, M. (2022b). The fossil record of crocodylians (Reptilia: Crocodylia) in Greece. In E. Vlachos (Ed.), Fossil vertebrates of Greece Vol. 1 ‐ basal vertebrates, amphibians, reptiles, Afrotherians, Glires, and primates (pp. 237–243). Springer Nature. https://doi.org/10.1007/978-3-030-68398-6_8.
Georgalis, G. L., Mennecart, B., & Smith, K. T. (2023). First fossil record of Varanus (Reptilia, Squamata) from Switzerland and the earliest occurrences of the genus in Europe. Swiss Journal of Geosciences, 116, 9. https://doi.org/10.1186/s00015-023-00440-5.
Georgalis, G. L., Rage, J.‐C., de Bonis, L., & Koufos, G. D. (2018). Lizards and snakes from the late Miocene hominoid locality of Ravin de la Pluie (Axios Valley, Greece). Swiss Journal of Geosciences, 111(1–2), 169–181. https://doi.org/10.1007/s00015-017-0291-2.
Georgalis, G. L., Szyndlar, Z., Kear, B. P., & Delfino, M. (2016). New material of Laophis crotaloides, an enigmatic giant snake from Greece, with an overview of the largest fossil European vipers. Swiss Journal of Geosciences, 109(1), 103–116. https://doi.org/10.1007/s00015-016-0210-y.
Georgalis, G. L., Villa, A., & Delfino, M. (2017). The last European varanid: demise and extinction of monitor lizards (Squamata, Varanidae) from Europe. Journal of Vertebrate Paleontology, 37(2), e1301946.
Georgalis, G. L., Villa, A., Ivanov, M., & Delfino, M. (2024). New diverse amphibian and reptile assemblages from the late Neogene of northern Greece provide novel insights into the emergence of extant herpetofaunas of the southern Balkans. Swiss Journal of Palaeontology, 143, 34. https://doi.org/10.1186/s13358-024-00332-7.
Georgalis, G. L., Villa, A., Ivanov, M., Vasilyan, D., & Delfino, M. (2019). Fossil amphibians and reptiles from the Neogene locality of Maramena (Greece), the most diverse European herpetofauna at the Miocene/Pliocene transition boundary. Palaeontologia Electronica, 22(3), 68. https://doi.org/10.26879/908.
Gray, J. E. (1827). A synopsis of the genera of saurian reptiles, in which some new genera are indicated, and the others reviewed by actual examination. The Philosophical Magazine, or Annals of Chemistry, Mathematics, Astronomy, Natural History, and General Science, 2(2), 54–58.
Hardwicke, T., & Gray, J. E. (1827). A synopsis of the species of saurian reptiles, collected in India by major‐general Hardwicke. The Zoological Journal, 3, 214–229.
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.
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. https://doi.org/10.1111/j.1475-4983.2010.00994.x.
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(9), 767–797. https://doi.org/10.1080/14772019.2017.1355338.
Kalaitzi, C., & Kostopoulos, D. S. (2025). Craniodental anatomy of late Ruscinian Trischizolagus (Leporidae: Lagomorpha) from Megalo Emvolon (Thessaloniki, Greece). Zoological Journal of the Linnean Society, 204(2), zlaf067. https://doi.org/10.1093/zoolinnean/zlaf067.
Kearney, M., Rieppel, O., & Wood, R. (2006). An investigation into the occurrence of Plicidentine in the teeth of Squamate reptiles. Copeia, 2006(3), 337–350.
Koufos, G. D., Syrides, G. E., & Koliadimou, K. K. (1991). A Pliocene primate from Macedonia (Greece). Journal of Human Evolution, 21(4), 283–294.
Laurenti, J. N. (1768). Specimen medicum, exhibens synopsin reptilium emendatam cum experimentis circa venena et antidota reptilium austracorum, quod authoritate et consensu. Joan Thomae.
Lazaridis, G. (2015). Study of the late Miocene vertebrate locality of Kryopigi and other localities of Kassandra peninsula, Chalkidiki (Greece). Systematics, taphonomy, paleoecology, biochronology. Ph.D. dissertation. Thessaloniki: Aristotle University of Thessaloniki.
Leidy, J. (1870). Descriptions of Emys jeanesi, E. haydeni, Baena arenosa, and Saniwa ensidens. Proceedings of the Academy of Natural Science of Philadelphia, 1870, 123–124.
Linnaeus, C. (1758). In R. Decima (Ed.), Systema Naturae, per regna Tria Naturae, secundum classes, ordines, genera, species, cum Characteri‐bus, Differentiis, Synonymis, Locis. Tomus I (10th ed.). Laurentii Salvii.
Merrem, B. (1820). Versuch eines systems der Amphibien (Vol. 8). J.C. Krieger.
Mertens, R. (1942a). Die Familie der Warane (Varanidae). Erster Teil: Allgemeines. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 462, 1–116.
Mertens, R. (1942b). Die Familie der Warane (Varanidae). Zweiter Teil: Der Schädel. Abhandlungen der Senckenbergischen Naturforschenden Gesellschaft, 465, 117–234.
Nopcsa, F. (1908). Zur Kenntnis der fossilen Eidechsen. Beitrage Zur Paläontologie Und Geologie Österreich‐Ungarns Und Des Orients, 21, 33–62.
Oppel, M. (1811). Die Ordnungen, Familien und Gattungen der Reptilien als Prodrom einer Naturgeschichte derselben. Joseph Lindauer.
Ouwens, P. A. (1912). On a large Varanus species from the Island of komodo. Bulletin du Jardin Botanique de Buitenzorg, Series, 2(1–6) 1911–1912, 1–3.
Owen, R. (1857). On the fossil vertebrae of a serpent (Laophis crotaloïdes, ow.) discovered by Capt. Spratt, R.N., in a tertiary formation at Salonica. Quarterly Journal of the Geological Society of London, 13, 196–199.
Pianka, E. R., King, D. R., & King, R. A. (Eds.). (2004). Varanoid lizards of the world. Indiana University Press. https://doi.org/10.2307/j.ctt2005wjp.
Rage, J.‐C. (2013). Mesozoic and Cenozoic squamates of Europe. Palaeobiodiversity and Palaeoenvironments, 93, 517–534. https://doi.org/10.1007/s12549-013-0124-x.
Ratnikov, V. Y. (2002). Late Cenozoic amphibians and squamate reptiles of the east‐European plain. Voronezh State University.
Rieppel, O. (1979). A functional interpretation of the varanid dentition (Reptilia, Lacertilia, Varanidae). Gegenbaurs Morphologisches Jahrbuch, 125(6), 797–817.
Sanz, J. (1977). Presencia de Varanus (Sauria, Reptilia) en el Plioceno de Layna (Soria). Trabajos Sobre Neogeno‐Cuaternario, 8, 113–125.
Schlegel, H. (1839). Abbildungen neuer oder unvollständig bekannter Amphibien, nach der Natur oder dem Leben entworfen und mit einem erläuternden Texte begleitet. Hermann.
Sion, G., & D'Amore, D. C. (2025). Varanid teeth asymmetry and correlation to body size. Journal of Developmental Biology, 13(1), 9. https://doi.org/10.3390/jdb13010009.
Smith, K. T., Bhullar, B.‐A. S., & Holroyd, P. A. (2008). Earliest African record of the Varanus stem‐clade (Squamata: Varanidae) from the early Oligocene of Egypt. Journal of Vertebrate Paleontology, 28(3), 909–913.
Smith, K. T., Bhullar, B.‐A. S., Köhler, G., & Habersetzer, J. (2018). The only known jawed vertebrate with four eyes and the Bauplan of the pineal complex. Current Biology, 28, 1101–1107.
Syrides, G. (1990). Lithostratigraphic, biostratigraphic and palaeogeographic study of the neogene‐quaternary sedimentary deposits of Chalkidiki peninsula, Macedonia, Greece. Ph.D. dissertation, Thessaloniki: Aristotle University of Greece.
Villa, A., Abella, J., Alba, D., Almécija, S., Bolet, A., Koufos, G., Knoll, F., Hernández, L. À., Morales, J., Robles, G. J., Sánchez, I., & Delfino, M. (2018). Revision of Varanus marathonensis (Squamata, Varanidae) based on historical and new material: Morphology, systematics, and paleobiogeography of the European monitor lizards. PLoS One, 13, e0207719. https://doi.org/10.1371/journal.pone.0207719.
Villa, A., Carnevale, G., Pavia, M., Rook, L., Sami, M., Szyndlar, Z., & Delfino, M. (2021). An overview of the late Miocene vertebrates from the fissure fillings of Monticino quarry (Brisighella, Italy), with new data on non‐mammalian taxa. Rivista Italiana di Paleontologia e Stratigrafia, 127(2), 297–354.
Villa, A., & Delfino, M. (2019). Fossil lizards and worm lizards (Reptilia, Squamata) from the Neogene and quaternary of Europe: an overview. Swiss Journal of Palaeontology, 138(2), 177–211.
Vlachos, E. (2022). The fossil record of turtles and tortoises (Reptilia: Testudines) in Greece. In E. Vlachos (Ed.), The fossil vertebrates of Greece Vol. 1 ‐ basal vertebrates, amphibians, reptiles, afrotherians, Glires, and primates (pp. 245–281). Springer Nature.
Vlachos, E., Kotsakis, T., & Delfino, M. (2014). The chelonians from the latest Miocene‐earliest Pliocene localities of Allatini and Pylea (East Thessaloniki, Macedonia, Greece). Comptes Rendus Palevol, 14, 187–205.
Weithofer, A. (1888). Beiträge zur Kenntniss der Fauna von Pikermi bei Athen. Beiträge Zur Paläontologie Österreich‐Ungarns, 6, 225–292.
Zerova, G. A., & Chkhikvadze, V. M. (1986). Neogene Varanids of the USSR. In Z. Roček (Ed.), Studies in Herpetology (pp. 689–694). Charles University of Prague.
Contributed Indexing: Keywords: Europe; Ruscinian; anatomy; monitor lizards; systematics
Entry Date(s): Date Created: 20251015 Date Completed: 20260703 Latest Revision: 20260726
Update Code: 20260726
PubMed Central ID: PMC13331538
DOI: 10.1002/ar.70068
PMID: 41090542
Βάση Δεδομένων: MEDLINE
Περιγραφή
ISSN:1932-8494
DOI:10.1002/ar.70068