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126 to 11.7 thousand years ago, Greece formed part of a dynamic Mediterranean landscape shaped by glacial cycles, flutuating sea levels, and biogenographic isolation. Within this setting, testudines were among the most resilient reptilian groups. Who occupied terrestrial, freshwater and occaisonally coastal habitat niches.

Greece preserves one of the richest fossil records of tortoises and turtles in the eastern Mediterranean. There are remains from the miocene to the holocene, representing multiple ecological strategies

Crete had endemic island tortoises like the testudo marginata cretensis (now extinct). It was larger than mainland relatives, and exhibited distinct shape and morphology. It was known from cave and fissure deposits across the island (Geranai IV, Zourida Cave, Simonelli Cave, Rethymnon fissures, and Sitia (open air)).

The tortoises here illustrate classic island evolutionary patterns. Gigantism, or reduced predation pressure, allowing larger body sizes. And, endism or long term isolated leading to distinct local forms and habitat flexibility. These tortoises lived alongside ofther endemic megafauna like dwarf elephants and deer too. Forming a uniquely insular ecosystem. Many thousands of years ago.

Testudo marginata cretensis likely disapeared because of climate fluctuations, and due to glacial, interglacial transitions; habitat shifts and possible early human pressures.

More north, in the Thermaikos gulf region excavations have revealed giant terrestrial tortoises (testudinidae); and small tortoises. Including earlier forms of chersine (testudu) hermanni; and, freshwater turtles, the genus mauremy. This region may represent the oldest hermanns tortoise lineage relevant to the pleistocene.

During the pleistocene, across Greece, the main fossil record shows at least 14 species across several famalies, and the dominance of testudinidae (tortoises). There were also freshwater turtles (geomydidae, emydidae), softshell turtles (trionychidae, and earlier periods). It likely included testudo marginata (marginated tortoise), testudo freca (greek tortoise), testudo hermanni (hermanns tortoise and linaege), and freshwater taxa such as mauremys.

Mediterranean shrubland, open forests, and river systems created habitat for these species. And, it allowed testudines to adapt to seasonal droughts, and temperature changes locally. It was a glacial refugium too, providing milder climates than northern Europe. More stable habitats, and a recolonization source for post glacial conditions.

Their characterizations, morphology and adaptations show they played an important role. Like domed or flared carapaces, especially on marginated tortoises. Hinged plastrons (allowing shell closure); and, larger size variations on Crete. It contributed to ecological survival as herbivorus grazers and browsers, seed dispersers, and prey for predators (on mainland).

By the late pleistocene, hominins (neanderthals, and early homo sapeins) were present. The testudines provided reliable food sources, and occasional raw materials.

Crete may have produced some larger testudine forms, like testudo marginata cretensis, and it is a phenomenon dating to pleistocene periods. The Thermaikos gulf provided a window into earlier ecosystems too. And, the mainland provided a persistant refugium through climate changes. It was a complex interplay between geography, climate and evolution. Together, these regions demonstrate testudines were not only survivors, but key ecological components of mediterranean ecosystems. Bridging deep evolutionary history with more modern fauna. Check it out today.

Bibliography: Vlachos, Evangelos. (2022). The fossil record of turtles from the Pleistocene of Crete (Greece). Comptes Rendus Palevol. 21. 771-799. 10.5852/cr-palevol2022v21a35

Garcia G, Pinton A, Valentin X, Kostopoulos DS, Merceron G, de Bonis L, Koufos GD. The earliest known crown-Testudo tortoise from the late Miocene (Vallesian, 9 Ma) of Greece. PLoS One. 2020 Apr 8;15(4):e0224783. doi: 10.1371/journal.pone.0224783. PMID: 32267839; PMCID: PMC7141670.

Evangelos Vlachos, Adán Pérez-García, Socrates Roussiakis, Georgios L. Georgalis, Benjamin P. Kear “Late Miocene Tortoises from Samos, Greece: Implications for European Neogene Testudinid Systematics and Distributions,” Journal of Vertebrate Paleontology, 39(6), (14 July 2020)

Georgios L. Georgalis, Benjamin P. Kear, The fossil turtles of Greece: An overview of taxonomy and distribution, Geobios, Volume 46, Issue 4, 2013, Pages 299-311, ISSN 0016-6995, https://doi.org/10.1016/j.geobios.2013.05.001. (https://www.sciencedirect.com/science/article/pii/S0016699513000430)

Georgios L. Georgalis, Andrea Villa, Evangelos Vlachos, Massimo Delfino, Fossil amphibians and reptiles from Plakias, Crete: A glimpse into the earliest late Miocene herpetofaunas of southeastern Europe, Geobios, Volume 49, Issue 6, 2016, Pages 433-444, ISSN 0016-6995, https://doi.org/10.1016/j.geobios.2016.09.004. (https://www.sciencedirect.com/science/article/pii/S0016699516300973)