Turtle Remains from the Late Miocene of the Cessaniti Area, Southern Italy—Insights for a Probable Tortonian Chelonian Dispersal from Europe to Africa Georgios L
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Georgalis et al. Swiss J Palaeontol (2020) 139:1 https://doi.org/10.1186/s13358-020-00202-y Swiss Journal of Palaeontology RESEARCH ARTICLE Open Access Turtle remains from the late Miocene of the Cessaniti area, southern Italy—insights for a probable Tortonian chelonian dispersal from Europe to Africa Georgios L. Georgalis1*, Gianni Insacco2, Lorenzo Rook3, Filippo Spadola2,4 and Massimo Delfno1,5 Abstract We here describe turtle remains from the late Miocene (Tortonian) of Cessaniti (Calabria, southern Italy), an area that recently has been palaeogeographically reconstructed as being, at that time of the Neogene, directly connected (or at least rather proximate) to northern Africa, instead of Europe. The material pertains to three diferent turtle clades, i.e., pan-trionychids, pan-cheloniids, and pan-geoemydids. Although the material is incomplete, it nevertheless permits a more precise identifcation for the pan-trionychid specimens, which are referred to the species Trionyx pliocenicus, as well as the pan-geoemydid, which is attributed to the genus Mauremys. Especially for the case of T. pliocenicus, the new Cessaniti specimens expand its geographic and stratigraphic distribution and further comprise the sole exist- ing material known for this species, considering that its holotype and so far only known material is currently lost. Overall, besides its taxonomic signifcance, the Cessaniti chelonian assemblage afords the potential for important biogeographic implications, attesting that the lineages of Trionyx and Mauremys could have potentially used the Sicily–Calabria arch for their dispersal from Europe to Africa during the Tortonian. The new turtle specimens further complement the associated mammal remains in envisaging the Cessaniti assemblage as a mosaic of both African and Eurasian (Pikermian) faunal elements. Keywords: Testudines, Trionyx, Mauremys, Pan-Cheloniidae, Miocene, Biogeography Introduction dramatically palaeonvironments and faunal composition Te palaeogeography of Europe and adjacent areas has (e.g., Rögl 1999; Delfno et al. 2003, 2013a; Koufos et al. been subjected to signifcant alterations throughout the 2003; Rook et al. 2006, 2008; Rage 2013; Georgalis et al. Neogene (Rögl 1999; Meulenkamp and Sissingh 2003). On 2016a, 2019a, b). Two such, palaeogeography-driven, Neo- certain occasions, drastic palaeogeographic changes have gene dispersal events are mostly known and have been facilitated waves of dispersals of vertebrate taxa from Africa more meticulously studied: one during the early Miocene to Europe during the Neogene, afecting tremendously the (Burdigalian), when the Afro-Arabian plate collided with faunal composition of both continents, with new incom- Eurasia, allowing direct terrestrial connection among the ing immigrants drastically outcompeting autochthonous two areas on the east via the so-called Gomphotherium animals, occupying novel ecological niches, and reshaping land bridge (Koufos et al. 2003, 2005; Georgalis et al. 2016a, 2019a) and another one during the latest Miocene (Messin- Editorial handling: D. Marty. ian) that was facilitated by the Messinian Salinity Crisis, *Correspondence: [email protected] when big parts of the Mediterranean Sea dried out (Hsü 1 Dipartimento di Scienze della Terra, Università di Torino, Via Valperga 1972; Torres-Roig et al. 2017; Georgalis et al. 2019b; see Caluso 35, 10125 Turin, Italy Full list of author information is available at the end of the article Carnevale et al. 2019, for a recent overview of the Italian © The Author(s) 2020. This article is licensed under a Creative Commons Attribution 4.0 International License, which permits use, sharing, adaptation, distribution and reproduction in any medium or format, as long as you give appropriate credit to the original author(s) and the source, provide a link to the Creative Commons licence, and indicate if changes were made. The images or other third party material in this article are included in the article’s Creative Commons licence, unless indicated otherwise in a credit line to the material. If material is not included in the article’s Creative Commons licence and your intended use is not permitted by statutory regulation or exceeds the permitted use, you will need to obtain permission directly from the copyright holder. To view a copy of this licence, visit http://creat iveco mmons .org/licen ses/by/4.0/. 1 Page 2 of 13 G. L. Georgalis et al. fossil record of the Messinian Salinity Crisis). Moreover, pan-trionychid costal MSNC 4591 originate from Cava geological data and the fossil record also show diferent Franzè. Part of the herein described material has been palaeogeographic reconstructions for the Aegean region, presented in the unpublished theses of Delfno (2002) and where the Greek mainland, islands (including also Crete), Chesi (2009). Comparative material of extinct and extant and Anatolia were united during the early Miocene (Lym- turtles belongs to the following collections: MDHC, berakis and Poulakakis 2010; Georgalis et al. 2016b), as well MGP-PD, MGPT-PU, MNHN, NHMW, and NHMUK. as the indication of unique insular environments (“palae- Authorships of turtle clades follow Joyce et al. (2004). oislands”) in the Gargano and Scontrone areas (Freudenthal 1971, 1972; Rook et al. 2006, 2008; Freudenthal and Martín- Geological and palaeoecological settings Suárez 2010), as well as in the Tuscany area (Rook et al. Te Cessaniti fossiliferous area consists of diferent local- 2006, 2008; Rook 2016), which are all now parts of the Ital- ities/quarries cropping out in the area between Capo ian mainland. Vaticano and Vibo Valentia on the Calabria Tyrrhenian Interestingly, fossil evidence from southern Italy has coast (Gramigna et al. 2008; Marra et al. 2017). Among recently allowed the identifcation of another, not yet so them, Cava Gentile has produced most of the vertebrate well-known palaeobioprovince testifed by sites in Calabria remains described so far (see Marra et al. 2017). Tis is and Sicily characterized by non-endemic mammals, coun- also the case with the turtle fnds that are presented in terparts of which were identifed in both North Africa and this paper (i.e., all turtle fossils originate from Cava Gen- Europe (Rook et al. 2008). Te identifcation of the elephan- tile, except for the Mauremys specimen MSCN 4380 tid proboscidean Stegotetrabelodon syrticus (which is of and the Trionyx costal MSNC 4591 that both come from clear African origin) in the late Miocene (Tortonian) of the Cava Franzè). Both Cava Gentile and Cava Franzè are Cessaniti area, Calabria, southern Italy, has led to the sug- considered to be of Tortonian age (Carone and Domn- gestion that, during that time interval, the Sicily–Calabria ing 2007; Marra et al. 2017; Stara and Marini 2018). Sev- arch (Siculo–Calabrian arc) was united with northern eral studies have dealt with the geology of the localities Africa, forming in fact part of the latter continent and not of the Cessaniti area during the past decades (Nicotera Europe (Ferretti et al. 2003). Further fnds of terrestrial 1959; Ogniben 1973; Moncharmont Zei and Montchar- mammals of Eurasian afnities in Cessaniti were envisaged mont 1987; Carone and Domning 2007; Gramigna et al. to denote an expansion of these faunal elements towards 2008; Marra et al. 2017). Te most recent review of geol- Africa (Marra et al. 2011). In order to improve the knowl- ogy, palaeoenvironmental, and stratigraphic setting of the edge of this palaeobioprovince, that has been based so far area is ofered by Marra et al. (2017), where the age range on mammals only, we here describe for the frst time rep- of Cava Gentile, on the basis of magnetostratigraphic and tile fnds from the Cessaniti area. Tis material, represented biostratigraphic control, has been suggested to be within by turtle remains recovered from those localities, pertains the range of 8.1 and 7.2 Ma (Marra et al. 2017). to three diferent lineages and provides an opportunity to Te Cava Gentile quarry of Cessaniti has yielded an assess the previous palaeogeographic reconstructions and array of marine invertebrate taxa (e.g., Gramigna et al. test the previously suggested biogeographic scenarios. 2008; Stara and Marini 2018), however, mammal fnds are Institutional abbreviations MDHC, Massimo Del- also known. Mammal fnds comprise marine taxa such as fno Herpetological Collection, University of Torino, the sirenian Metaxytherium and cetaceans (Neviani 1886; Italy; MGP-PD Museo di Geologia e Paleontologia Carone and Domning 2007; Marra et al. 2016), but also dell’Università di Padova, Padua, Italy; MGPT-PU Museo terrestrial forms such as the proboscidean S. syrticus, the di Geologia e Paleontologia, Università degli Studi di endemic rhinocerotid Ceratotherium advenientis, the Torino, Turin, Italy; MNHN, Muséum national d’Histoire girafds Bohlinia and Samotherium, the bovid Trago- naturelle, Paris, France; MSNC, Museo Civico di Storia portax cf. rugosifrons, while (as yet undescribed) reports Naturale di Comiso, Comiso, Italy; NHMW, Naturhis- of smaller bovids and hippopotamids also exist (Ferretti torisches Museum Wien, Vienna, Austria; NHMUK, Nat- et al. 2003; Marra et al. 2011; Marra 2018; Pandolf et al. ural History Museum, London, United Kingdom. 2019). Fossil and sedimentological evidence (Marra et al. 2017) suggests that the palaeoenvironment of the Ces- Materials and methods saniti area was a mosaic environment with open spaces, Te turtle material described herein is curated at MSNC. probably similar to the modern savannah,