Geobios xxx (xxxx) xxx Contents lists available at ScienceDirect Geobios journal homepage: www.elsevier.com/locate/geobio Original article The Scontrone turtles – A new insular testudinoid fauna from the late Miocene of the Central Mediterranean q Georgios L. Georgalis a,⇑, Massimo Delfino b,c,⇑ a Palaeontological Institute and Museum, University of Zurich, Karl Schmid-Strasse 4, 8006 Zurich, Switzerland b Dipartimento di Scienze della Terra, Università di Torino, 35, via Valperga Caluso, 10125 Torino, Italy c Institut Català de Paleontologia Miquel Crusafont, Universitat Autònoma de Barcelona, Edifici ICTA -ICP, Carrer de les Columnes s/n, Campus de la UAB, 08193 Cerdanyola del Vallès, Barcelona, Spain a r t i c l e i n f o a b s t r a c t Article history: We here describe a small turtle assemblage originating from the early Tortonian (late Miocene) palaeois- Received 2 January 2021 land of Scontrone, central Italy, a locality previously known mostly for its endemic mammals and giant Accepted 15 May 2021 birds, which were otherwise shared only with the Gargano localities, another fossiliferous area belonging Available online xxxx to the same palaeobioprovince. The fossil turtle remains from Scontrone are referred to the geoemydid Mauremys sp. and a so far unidentified large-sized testudinid. The biogeographic origins of the Keywords: Scontrone insular chelonians are discussed. The Scontrone geoemydid adds to the known occurrences Testudines of Mauremys in the late Miocene of the Mediterranean. The Scontrone large tortoise represents the oldest Geoemydidae Testudinidae known Mediterranean insular testudinid, predating significantly the well-known Quaternary endemic Palaeoisland island tortoises of the area. Biogeography Ó 2021 The Author(s). Published by Elsevier Masson SAS. This is an open access article under the CC BY-NC- Miocene ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/). Introduction periods of time (Patacca et al., 2008a, 2008b, 2013). These two localities have been grouped in the same palaeobioprovince, the Mediterranean islands offer a unique insight into the evolution Apulo-Abruzzi area (Rook et al., 2008). The majority of the different of several vertebrate lineages and witness the acquisition of fissures of Gargano pertain to a timespan between the late Miocene extreme and astonishing insular adaptations and species radiations and the Pliocene and have yielded a considerably rich fossil fauna (Van der Geer et al., 2010; Herridge and Lister, 2012; Palombo over the past decades (Freudenthal, 1971, 1972, 1976, 1985; et al., 2013; Georgalis et al., 2019). Especially for testudinid turtles, Ballmann, 1976; Leinders, 1984; Abbazzi et al., 1996; Delfino Mediterranean islands are long known to have drastically affected et al., 2007; Masini et al., 2010, 2013; Masini and Fanfani, 2013; the evolution of this lineage, witnessing the radiation and extinc- Freudenthal et al., 2013; Villier and Carnevale, 2013; Meijer, tion of an array of unique, endemic, and bizarre forms across differ- 2014; Van Der Geer, 2014; Savorelli and Masini, 2016; Pavia ent insular ecosystems (Leith Adams, 1877; Bate, 1914; Abbazzi et al., 2017; Pavia, 2013); younger, Early Pleistocene sites, known et al., 2008; Luján et al., 2017). as Pirro Nord are also known in Gargano (for the reptiles see, Besides the islands that are present today in the Mediterranean among others, Delfino and Bailon, 2000; Blain et al., 2019). Scon- Sea, other palaeoislands also existed in the area during its Cenozoic trone is older (Tortonian) and has yielded a poorer, though still past, that are currently, however, part of mainland Europe (Lanza, unique faunal assemblage, comprising also endemic mammals 1984). The most prominent and well known among such cases rep- and birds with peculiar morphologies (Rustioni et al., 1992; resent the famous fossiliferous localities in Gargano in southern Mazza and Rustioni, 1996, 2008, 2011; Savorelli et al., 2017; Italy and Scontrone in central Italy, which were during the late Meijer, 2014; Van Der Geer, 2014; Pavia et al., 2017). Miocene, parts of the Abruzzo-Apulia Platform that has been Turtle remains from these unique insular environments of the repeatedly isolated from neighbouring mainland areas for long late Neogene palaeoislands of Gargano and Scontrone have been only poorly known, being solely represented by preliminary descriptions in a thesis of scarce testudinid remains from Gargano q Corresponding editor: Pierre-Olivier Antoine. (Delfino, 2002) and simple mentions (Mazza and Rustioni, 2008; ⇑ Corresponding authors. Patacca et al., 2013; Collareta et al., 2020) or partial descriptions E-mail addresses: georgios.georgalis@pim.uzh.ch (G.L. Georgalis), massimo. in a thesis (Chesi, 2009b) of Mauremys from Scontrone. Here we delfino@unito.it (M. Delfino). https://doi.org/10.1016/j.geobios.2021.05.001 0016-6995/Ó 2021 The Author(s). Published by Elsevier Masson SAS. This is an open access article under the CC BY-NC-ND license (http://creativecommons.org/licenses/by-nc-nd/4.0/). Please cite this article as: G.L. Georgalis and M. Delfino, The Scontrone turtles – A new insular testudinoid fauna from the late Miocene of the Central Mediterranean, Geobios, https://doi.org/10.1016/j.geobios.2021.05.001 G.L. Georgalis and M. Delfino Geobios xxx (xxxx) xxx formally describe for the first time all the chelonian remains col- Material and methods lected so far (a few specimens are exposed on the rock surface of the site) from the palaeoisland of Scontrone, which document the All material described herein is permanently curated at the col- presence of at least two non-marine turtle lineages. The potential lections of the Centro di Documentazione Paleontologico ‘‘Hoplito- affinities and biogeographic origins of the new turtles from the meryx”, Scontrone, Italy. Taxonomy and nomenclature of turtle Scontrone palaeoisland are discussed. clades follows Joyce et al. (2020), Joyce et al. (2021). Institutional abbreviation: SCT, Centro di Documentazione Geological settings Paleontologico ‘‘Hoplitomeryx”, Scontrone, Italy. The fossiliferous locality of Scontrone is situated in the Parco Systematic palaeontology Nazionale d’Abruzzo, Lazio e Molise, in Abruzzo region, Central Italy (41°450 15.5400 N, 14°020 13.1400 E; Fig. 1). The Scontrone fossil Testudines Batsch, 1788 [Joyce et al., 2020] site consists of tidal-flat aeolian calcarenites at the base of the Pan-Testudinoidea Joyce, Parham, and Gauthier, 2004 [Joyce Lithothamnion Limestone, a Miocene carbonate ramp widespread et al., 2021] in the central-southern Apennines (Savorelli et al., 2017). More Testudinoidea Fitzinger, 1826 [Joyce et al., 2021] details about the geology of the locality can be found in Patacca Geoemydidae Theobald, 1868 [Joyce et al., 2021] et al. (2008a), Patacca et al. (2013). Genus Mauremys Gray, 1869 The locality is ubiquitously considered to pertain to a palaeois- Mauremys sp. land. The age of Scontrone is ascribed to the early Tortonian stage Figs. 2, 3 of the late Miocene, ca. 9 Ma (Patacca et al., 2013; Savorelli et al., Material: SCT 371a (a neural), SCT 371b (two costals and nine 2017). Associated mammal fauna of Scontrone includes erinaceids, shell fragments); SCT 26a (a costal); SCT 26b (a costal); rodents, and hoplitomerycids (Mazza and Rustioni, 1996, 2008, SCT 127a (a fragmentary costal); SCT 43 (five costal fragments 2011; Savorelli et al., 2007; Van Der Geer, 2014), while giant and a bridge peripheral); SCT 256 (a peripheral); SCT 285 (a non-flying birds are also present (Pavia et al. 2017). Among rep- peripheral); SCT 258 (a peripheral); SCT 421 (a peripheral); tiles, only a crocodylian (cf. Crocodylus sp.) has been described so SCT 416 (a peripheral); SCT 463 (a peripheral); SCT 218 (three far from Scontrone (Delfino and Rossi, 2013). peripherals); SCT 422 (a bridge peripheral); SCT 423 (fragment of a bridge peripheral); SCT 140a (a fragmentary ?suprapygal); SCT 233 (right epiplastron); SCT 214b (a left epiplastron); SCT 251 (a right epiplastron); SCT 334 (a right epiplastron frag- ment); SCT 253 (a fragmentary entoplastron); SCT 301 (a right hyoplastron); SCT 535 (a right hyoplastron on a slab); SCT 249 (a left hyoplastron); SCT 248 (a fragment of hyoplastron or hypoplastron); SCT 339 (a fragment of hyoplastron or hypoplas- tron); SCT 396 (a fragment of hyoplastron or hypoplastron); SCT 270 (a right hypoplastron); SCT 310 (a partial left hypoplas- tron); SCT 358 (a fragmentary hyo-hypoplastron); SCT 349 (a right xiphiplastron on a slab); SCT 239 (a left xiphiplastron); SCT 259 (several shell fragments on matrix); SCT 119 (an ilium). Description Carapace: The available carapace material consists of a neural, several costals and peripherals, and a probable suprapygal (Fig. 2). The single available neural is SCT 371a (Fig. 2(A)); it is not complete, as its anterolateral edge is broken off, but it is clear that the original general shape was hexagonal, with a concave anterior edge, and posterolateral edges much longer than the anterolateral ones. Most costals are fragmentary (Fig. 2(B, C)). Costal SCT 127a shows the possible trace of a long plastral buttress developed across a few centimetres along its ventral surface. Most shell remains available are peripherals, including bridge ones. SCT 421 is a peripheral pertaining to the beginning or the end of the bridge. Its external surface is characterized by a pleuro- marginal sulcus situated far from the costo-peripheral suture; it bears an intermarginal sulcus. SCT 416 is a complete peripheral originating from the anterior or posterior portion of the carapace, showing a clear intermarginal sulcus, but not a pleuro-marginal one. SCT 463 is a large-sized peripheral (probably peripheral I) that clearly shows the presence of a pleuro-marginal sulcus, as well as an intermarginal one (Fig. 2(F, G)). The peripherals of the bridge are relatively thin if compared with most of the plastral elements. SCT 218 consists of three peripherals of which the central one Fig. 1. The late Miocene (Tortonian) fossiliferous locality in Scontrone, Abruzzo, (which is also the most complete) seems to pertain to the left front Italy. A. General panoramic view with the site (covered by a sliding protection) in of the bridge (Fig. 2(D, E)). It possesses pleuro-marginal and inter- the foreground. B. Close up with fossils still embedded in the rock. marginal furrows; prior to the beginning of the bridge cavity, at its 2 G.L. Georgalis and M. Delfino Geobios xxx (xxxx) xxx Fig. 2. Carapace elements of Mauremys sp. from the late Miocene (Tortonian) of Scontrone, Abruzzo, Italy. A. Neural SCT 371a in dorsal view. B. Costal SCT 26a in dorsal view. C. Costal SCT 26b in dorsal view. D, E. Three left bridge peripherals SCT 218 in anteroventral (D) and anterodorsal (E) views. F, G. Peripheral SCT 463 in dorsal (F) and ventral (G) views. Scale bars: 10 mm. ventral surface, there is an elongated depression that could house the last (likely XII) marginals, whereas a transversal sulcus is inter- the musk duct foramen (musk pore). SCT 140a is a fragmentary preted as the sulcus between the last marginals and the fifth ver- element that could represent a suprapygal. It is arched in cross- tebral scute. section, with sutures preserved on its ?posterior and ?left sides. Plastron: The plastral anatomy is relatively better known than A visible longitudinal sulcus is interpreted as the sulcus between that of the carapace, with plastron represented by epiplastra, an 3 G.L. Georgalis and M. Delfino Geobios xxx (xxxx) xxx Fig. 3. Plastron of Mauremys sp. from the late Miocene (Tortonian) of Scontrone, Abruzzo, Italy. A–D. Left epiplastron SCT 214b in dorsal (A), ventral (B), drawing of dorsal (C), and drawing of ventral (D) views. E–I. Right epiplastron SCT 233 in ventral (E), dorsal (F), anterior (G), drawing of ventral (H), and drawing of dorsal (I) views. J, K. Right hyoplastron SCT 535 on slab (J) and its drawing (K). L, M. Right xiphiplastron SCT 349 on slab, in lateral (L) and dorsal (M) views. N, O. Left xiphiplastron SCT 239 in ventral (N) and drawing of ventral (O) view. Scale bars: 10 mm. entoplastron, hyo-hypoplastra, and xiphiplastra (Fig. 3). SCT 214b to a distinct notch on the epiplastron (corresponding also to the is a left epiplastron characterized by a slightly developed epiplas- lateral edge of the epiplastral lip). The gularo-humeral sulcus inter- tral lip (Fig. 3(A–D)). The latter is medially concave (well visible sects the epi-entoplastral suture ca. 7 mm from the inter- in anterior view). The gularo-humeral sulcus laterally corresponds epiplastral suture indicating that the gulars significantly pene- 4 G.L. Georgalis and M. Delfino Geobios xxx (xxxx) xxx trated the entoplastron. There is no trace of a gular pocket, but the deep anal notch of the xiphiplastra, suggest a referral to rather only a weak depression posterior to the gular. SCT 251 is the genus Mauremys (Hervet, 2000; Claude et al., 2007). an incomplete right epiplastron, with an overall morphology con- gruent with that of SCT 214b, but slightly larger and thicker, being approximately 46 mm wide (due to erosion, this mediolateral width could be slightly larger in life). SCT 233 and SCT 334 corre- Pan-Testudinidae Joyce, Parham, and Gauthier, 2004 [Joyce spond to right epiplastra (Fig. 3(E–I)). SCT 253 is a fragmentary et al., 2021] entoplastron. Due to its fragmentary status, it is difficult to assess Testudinidae Gray, 1825 [Joyce et al., 2021] its exact shape, but it seems that it was wider than long. The sulci Testudinidae indet. (large form) visible on the external surface clearly indicate that the gulars cov- Figs. 4–7 ered the anterior tip of the entoplastron and that the pectorals cov- Material: SCT 118 (a partial posterior portion of a plastron ered its posterior portion (i.e., the humero-pectoral sulcus crosses partly preserved on a slab); SCT 127b–d (three plastral frag- the entoplastron). On its internal surface there is a clearly elon- ments); SCT 216a (a large shell fragment/unidentified laminar gated xiphoid scar. SCT 301 is a fragmentary right hyoplastron, bone); SCT 237 (a ?pectoral girdle on a slab); SCT 163 (a pec- partly covered by matrix (especially at the sutures with the epi- toral girdle); SCT 203 (a fragment of a ?pubis); SCT 308 (a plastron and entoplastron). Its anteroposterior length is ca. humerus); SCT 287 (a fragment of an ilium); SCT 236 (a frag- 67 mm long, whereas its maximum thickness reaches the ment of a ?humerus); SCT 64 (a caudal vertebra). 9.7 mm. This specimen preserves the humero-pectoral and Description pectoro-abdominal sulci. The position of the humero-pectoral sul- cus clearly indicates that it crossed the entoplastron quite far from Plastron: No carapace material is present and therefore our its posterior edge. The humero-pectoral sulcus is anteriorly con- knowledge on the shell anatomy of this taxon is based exclusively cave and laterally terminates in correspondence at the weak, but on its plastral remains (Figs. 4, 5). SCT 118 preserves the right still evident, notch in the lateral side of the anterior lobe of the hypoplastron and xiphiplastron (Fig. 4). Most of its external surface plastron. The pectoro-abdominal sulcus is anteriorly convex and is covered by sediment but the exposed areas are rugose due to a medially terminates far from the hyo-hypoplastral suture; its lat- prominent porosity (in the posterior tip of the xiphiplastron) or eral termination cannot be evaluated due to preservational rea- an irregular net of fine grooves (in the anterolateral sector of the sons. SCT 535 is a right hyoplastron on a slab (Fig. 3(J, K)). The hypoplastron). On the exposed area there is no evidence of sulci humero-pectoral sulcus is clearly concave anteriorly. The left by the scutes. The internal surface of both elements is rela- pectoro-abdominal sulcus is equally undulating but is convex ante- tively smooth. In the hypoplastron, its posterior edge and part of riorly in the central and middle portions of the hyoplastron, while its posterolateral area are characterized by a long and nearly hor- it forms a deep anterior concavity close to its lateral edge. SCT 249 izontal lateral buttress. The lateral edges of both the hypoplastron is a left hyoplastron whose external surface is so damaged that no and xiphiplastron is not preserved and therefore, the original shape sulci are visible. SCT 310 is a left hypoplastron that is only partially of the lateral edge of the posterior plastral lobe cannot be assessed. preserved. On its external surface, the abdomino-femoral forms an The xiphiplastron is nearly complete. This right xiphiplastron is anterior convexity close to the lateral edge of the element. SCT 310 rather thin (3 mm) along its sagittal suture with the (missing) left is slightly thickened (13.7 mm instead of 9.4 mm) at around its xiphiplastron but is comparatively much thicker at its posterior medial suture, in correspondence to the dorsal projection for the end, where a sort of ‘‘pad” nearly reaches 30 mm of thickness bridge buttress. Some further fragmentary elements are partly (the presence of fractures hinders a precise measurement of its embedded in matrix but a definite identification cannot be made, thickness). Both the dorsal (visceral) and the ventral (external) sur- such as the possible right hypoplastron SCT 270 and several frag- faces of this posterior ‘‘pad” are distinctly convex, but it is not clear ments of hyoplastra or hypoplastra (SCT 248, SCT 339, and whether the ventral convexity reflects the original morphology or SCT 396) that show only their visceral surfaces. SCT 349 is a right whether it is due to preservational reasons. However, this convex- xiphiplastron on a slab, situated along with an unidentified bone ity, along with the general structure of the xiphiplastron and that (Fig. 3(L, M)). This element shows only the visceral surface. Its lat- of the hypoplastron, impart to the posterior lobe of the plastron eral edge has a modest notch, corresponding to the femoro-anal a marked ventral concavity that possibly extended on the anterior sulcus, and a deep anal notch. The posterolateral edge of the lobe as well. The sagittal suture of this right xiphiplastron with its xiphiplastron is rounded. The structure of the anterior edge of missing left counterpart is ca. 80 mm long; the suture with the the element indicates the presence of a suture (not of a hinge). hypoplastron is ca. 11 mm long, but it should have been slightly The thickness of the xiphiplastron, in the area close to its anterior longer as both elements are laterally incomplete. Specimens suture, is 12 mm. SCT 239 is an incomplete left xiphiplastron, miss- SCT 127b–d seem also to represent plastral fragments (Fig. 5). ing its anterior portion (Fig. 3(N, O)). None of them shows any sulcus on the external surface. In these Appendicular skeleton: Only a single appendicular element is plastral fragments, the external surface possesses irregular furrows preserved, i.e., the ilium SCT 119. The glenoid cavity is not visible, (as in SCT 118), whereas the internal surface is rather smooth, with due to preservational reasons, but the area surrounding it is well the exception of possible vascular sulci. preserved. The distal (dorsal) edge is poorly preserved but it seems Appendicular skeleton: SCT 237 is a slab with a fragment of a that some residues of concretion are hosted in a lateral, elongated possible pectoral girdle, preserving the glenoid cavity and the base concavity. of both the scapula and acromion (Fig. 6(A)). The rugose area close to the glenoid cavity could represent the contact surface with the Remarks: The overall shape of the preserved skeletal elements coracoid. If so, the acromion is a slightly flattened structure that along with the probable presence of musk duct foramina is con- is a little thicker than the scapula. SCT 163 is a morphologically sistent with a referral of the material to Geoemydidae (Claude congruent element with SCT 237, but being isolated, it is more and Tong, 2004; Garbin et al., 2018; Joyce et al., 2021). Further- informative (Fig. 6(B)). Both the scapula and the acromion preserve more, the paired marginals XII entering a probable suprapygal, only the area close to the glenoid cavity, which is filled by matrix. the gulars reaching and overlapping the entoplastron, the The contact surface with the coracoid is located on the ventral half humero-pectoral sulcus crossing the entoplastron, as well as of the acromion, close to the glenoid cavity. This surface is sur- 5 G.L. Georgalis and M. Delfino Geobios xxx (xxxx) xxx Fig. 4. Plastron of Testudinidae indet. (large form) from the late Miocene (Tortonian) of Scontrone, Abruzzo, Italy. Right hypoplastron and xiphiplastron SCT 118 in dorsal (A), ventral (B), and posterolateral (C) views. Scale bar: 50 mm. mounted by a distinct tuberosity that is prolonged in a sort of ridge face opposite to the groove has, towards the distal epiphysis, a lon- along the dorsal edge of the acromion. Both the scapula and the gitudinal modest convexity delimited on both sides by two weak acromion are flattened and weakly concave on the surface (as concavities. SCT 236 is a rather fragmentary specimen that could the point where the contact surface is), but are slightly convex possibly also pertain to a humerus. on the other side. A marked concavity corresponds to the above SCT 287 could probably represent the dorsal epiphysis of a very described tuberosity on the other side of the bone. SCT 203 is a stout and large-sized right ilium (Fig. 7(C, D)). The epiphysis itself large element that could represent the proximal ramus of a partial is not complete and the posterodorsal process is better preserved pubis (Fig. 6(C, D)). It possesses a triradiate structure with a broad than the anterior one. The minimum diameters of the preserved and comparatively thick base distally, thinning into an arched front portion are 28 and 17 mm, respectively. laterally, which continues into two processes that are broken off. Vertebra: SCT 64 is a partially preserved and slightly deformed One of the two sides of the thick base is rounded, the other is caudal vertebra (centrum length = 26 mm) (Fig. 7(E, F)). The neural ridged. arch is severely damaged, though the elongate and slightly tilted SCT 308 is a fragmentary humerus preserving part of its distal prezygapophyseal facets are still visible. Part of the left postzy- portion, as the distal articulations are missing (Fig. 7(A, B)). The gapophyseal facet is also present. The left transverse process in maximum preserved length of the specimen is of 43 mm. The approximately horizontal (pointing ventrally only rather slightly) ectepicondylar groove for the radial nerve is well visible. The sur- and laterally directed. It is flattened but dorsally provided by a 6 G.L. Georgalis and M. Delfino Geobios xxx (xxxx) xxx Fig. 5. Plastron of Testudinidae indet. (large form) from the late Miocene (Tortonian) of Scontrone, Abruzzo, Italy. Plastral fragment SCT 127b in dorsal (A) and ventral (B) views. Scale bar: 20 mm. Fig. 6. Appendicular elements of Testudinidae indet. (large form) from the late Miocene of Scontrone, Abruzzo, Italy. A. ?Pectoral girdle SCT 237 on a slab. B. Pectoral girdle SCT 163. C, D. Partial ?pubis SCT 203 in dorsal (C) and ventral (D) views. Scale bars: 10 mm. modest keel. The neural canal is proportionally small. The vertebra Remarks: The presence of developed posterior buttresses on is procoelous and is characterized by two separated articular facets the plastron, along with the general shape of the elements both anteriorly and posteriorly. The cotyle faces anteroventrally, denote a referral to testudinids (Vlachos, 2018). More particu- whereas the condyle posterodorsally. The cotylar surfaces are sep- larly, within testudinids, the presence of such long and well- arated by an evident notch. The ventral surface of the centrum is developed buttresses in the posterior portion of the plastron flattened, though a slight longitudinal keel is present at its mid- would hint for affinities with geochelonans. Within geochelo- line. nans, prominent posterior buttresses are characteristic of the 7 G.L. Georgalis and M. Delfino Geobios xxx (xxxx) xxx Fig. 7. Appendicular and vertebral elements of Testudinidae indet. (large form) from the late Miocene of Scontrone, Abruzzo, Italy. A, B. Partial humerus SCT 308 in posterior (A) and anterior (B) views. C, D. Partial right ilium SCT 287 in anterior (C) and posterior (D) views. E, F. Caudal vertebra SCT 64 in left lateral (E) and ventral (F) views. Scale bars: 10 mm. European genus Titanochelon Pérez-García and Vlachos, 2014 any of the two testudinoid forms described above, though the (Pérez-García and Vlachos, 2014 (Pérez-García and Vlachos, presence of an additional taxon cannot be ruled out with cer- 2014; Vlachos and Rabi, 2018). However, more material is def- tainty. Nevertheless, affinities with trionychids, which are initely needed in order to conclude whether the Scontrone tor- otherwise present in the Tortonian of Italy (Georgalis and toise has affinities with Titanochelon or even with some African Joyce, 2017; Georgalis et al., 2020), can be readily discarded, form, such as Centrochelys Gray, 1872 (which also belongs to as the Scontrone shell fragments lack the characteristic sculp- geochelonans). In any case, the size of the preserved elements turing pattern of that turtle group. denotes that they pertain to a relatively large form with a rela- tively short posterior lobe of the plastron, though any estima- tions about its probable carapace or plastron length would be Discussion speculative. The Scontrone palaeoisland Although not so well known as the nearby Gargano palaeois- Testudines indet. land, Scontrone is characterized by an astonishing endemic fauna, Material: SCT 127e–h (four shell fragments), SCT 5, SCT 214c, comprising, among others, large erinaceids, the peculiar hoplito- SCT 214d, SCT 289, SCT 298, SCT 302, SCT 360, SCT 378, merycids, and large non-flying birds (Van Der Geer, 2014; Pavia SCT 472, SCT 474, SCT 408, SCT 485, SCT 497, SCT 498, et al., 2017; Savorelli et al., 2017). Several of these insular taxa have SCT 500, SCT 501, SCT 503, SCT 508, SCT 509, SCT 511, SCT 512 congeneric relatives only in Gargano and nowhere else in the (21 shell fragments), and SCT 106, SCT 387, and SCT 394 (three world, such as the iconic large erinaceid Deinogalerix Freudenthal, ?shell fragments). 1972 (Savorelli et al., 2017). Hoplitomerycids are also known Remarks. Besides the above described specimens, the assem- exclusively from Gargano and Scontrone: the presence of different blage comprises also several shell fragments that cannot be but still closely related genera in each palaeoisland, i.e., Hoplito- more precisely identified. Probably these could also pertain to meryx Leinders, 1984, in Gargano, and Scontromeryx Van Der Geer, 2014, in Scontrone (Leinders, 1984; Mazza and Rustioni, 8 G.L. Georgalis and M. Delfino Geobios xxx (xxxx) xxx 1996, 2011; Mazza, 2013; Van Der Geer, 2014) or of a single genus et al., 2019; Georgalis et al., 2020). As such, a direct terrestrial cor- (Hoplitomeryx) in both palaeoislands is still debated (Mazza et al., ridor would not be strictly necessary for these turtle taxa to reach 2016). The large non-flying anseriform bird Garganornis ballmanni Scontrone. Therefore, the new geoemydid and large testudinid Meijer, 2014, is also found exclusively in Gargano and Scontrone, from Scontrone could represent the product(s) of some dispersal notably being represented by conspecific finds in both palaeois- (s) during the late Paleogene and/or Neogene, though we have to lands (Meijer, 2014; Pavia et al., 2017). What appears rather leave open the possibility whether such dispersal(s) took place important is the fact that, unlike Gargano that represents a times- from either Africa, or the Balkans or Central Europe. Nevertheless, pan through part of the Neogene, the Scontrone palaeofauna is in particular for the geoemydid, a Paleogene or early Neogene Afri- strictly confined to the early Tortonian, offering thus a definite can origin seems unlikely, as this lineage apparently dispersed to stratigraphic accuracy and a unique glimpse into insular palaeoen- Africa from Europe only during the Tortonian (Georgalis et al., vironments of the Central Mediterranean area of that time. 2020) and is totally absent from pre-Tortonian sediments of The origins and exact timing of dispersal of the Scontrone faunal Afro-Arabia (Georgalis et al., 2021); therefore, if Mauremys reached assemblage have been a matter of debate, with different scenarios Scontrone following a southern to northern route, then this would variously proposed (e.g., Abbazzi et al., 1996; Mazza and Rustioni, not predate the Tortonian, implying that geoemydids were only 1996, 2008; Masini et al., 2002, 2008, 2013; Rook et al., 2006; new-comers on the palaeoisland, similarly to the case of their cro- Freudenthal and Martín-Suárez, 2010). In general, the whole codylian co-inhabitant (Delfino and Rossi, 2013). Abruzzo-Apulia Platform, to which Scontrone belonged, was iso- The identification of Mauremys in the Scontrone assemblage lated from Continental Europe at least since the early Oligocene adds to the known distribution of this genus in Italy, which con- and up to the latest Miocene (Mazza, 2013). In fact, it has been sug- sists of a ubiquitous faunal element throughout large part of the gested that the last interconnection of the Platform was through a Neogene and Quaternary, being nevertheless, currently extinct terrestrial land corridor in the Adriatic with northwestern Balkans from the area (Sacco, 1889; Portis, 1890, 1896; Kotsakis, 1981; during the early Oligocene, ca. 29 Ma (Patacca et al., 2008b; Mazza, Delfino and Bailon, 2000; Girotti et al., 2003; Chesi et al., 2007, 2013). It is thus not fully settled whether Gargano and Scontrone 2009a; Colombero et al., 2017; Collareta et al., 2020; Georgalis were colonized one or multiple times independently, though et al., 2020). It further attests for a wider distribution of Mauremys recent studies have provided strong evidence for the latter case, throughout the Mediterranean during the Tortonian (Georgalis i.e., multiphasic colonization events (Rook et al., 2006; Savorelli et al., 2016, 2020). As the present material cannot afford an identi- and Masini, 2016; Savorelli et al., 2017). In particular, different sce- fication to the species level, it cannot be determined whether the narios have been proposed for the most famous inhabitants of the Scontrone geoemydid represents an endemic insular form or Abruzzo-Apulia Platform: for Deinogalerix it has been suggested whether it is conspecific with any of the mainland European taxa, that its origins should be traced in an early to middle Miocene including the Italian extinct species, i.e., the late Miocene Maure- ancestor (Savorelli et al., 2017), whereas for the peculiar hoplito- mys campanii (Ristori, 1891), the Pliocene Mauremys portisii merycids that they arrived in the area already by the Oligocene (Sacco, 1889), and the Plio-Pleistocene Mauremys etrusca (Portis, (Mazza and Rustioni, 2011). 1890). As for the large-sized tortoise from Scontrone, its exact affinities Palaeobiogeography of the Scontrone turtles are also hindered by its poor preservational status, which mirrors the one of the few, likewise large, testudinid remains coming from Taking into consideration the complex biogeographic history of Gargano (Delfino, 2002). Large to giant tortoises were present in the insular Abruzzo-Apulian faunas discussed above, it seems a Europe throughout most of the Cenozoic (Lapparent de Broin, rather plausible scenario to adhere that the geoemydid and testu- 2002; Georgalis and Kear, 2013; Pérez-García and Vlachos, 2014; dinid ancestors of the Scontrone chelonians described herein are of Vlachos et al., 2020). Similarly, large-sized testudinids were also Balkan or even Western Asian origin that arrived in the area during present during the Cenozoic of northern Africa, though their fossil the Oligocene. For Mauremys, the possibility of an Oligocene dis- record there is significantly more poorly documented in compar- persal from the Balkans to Scontrone palaeoisland is hindered by ison with their European counterpart (Lapparent de Broin, 2002; the rather poor and limited fossil record of this genus in the Pale- Georgalis et al., 2021). In the Mediterranean Islands, testudinids ogene of Europe (Hervet, 2004), and particularly in the Balkan have a relatively rich fossil record, that comprises an array of ende- Peninsula. In fact, the only Oligocene taxon from the Balkans (but mic forms during the late Neogene and Quaternary of several also from the whole Eastern Europe) that could potentially pertain islands (e.g., Malta, Balearic Islands, Sardinia, Sicily), but all of to Mauremys is Mauremys strandi (Szalai, 1934) from the early Oli- these are from Plio-Pleistocene localities, thus postdating the Tor- gocene of Romania (Szalai, 1934; Młynarski and Mészáros, 1963), tonian age of the Scontrone material (Leith Adams, 1877; Bate, however, the generic placement of this species is still far from 1914; Delfino, 2002; Abbazzi et al., 2008; Luján et al., 2017). This clear. Also, two further Balkan taxa are Palaeochelys rueckerti Sch- diversity and abundance of Mediterranean insular testudinids, leich, 1994, and Palaeochelys turcica Schleich, 1994, both from the strongly evokes the situation in different oceanic islands across late Oligocene or early Miocene of the European part of Turkey the globe. There, several extant and recently (or at least relatively (Schleich, 1994), however, these are represented by rather incom- recently) extinct endemic testudinids flourished. These include the plete material and their affinities with Mauremys are not yet famous and diverse radiations in the Galapagos Islands (Chelonoidis resolved. Fitzinger, 1836; extant), the Mascarenes (Cylindraspis Fitzinger, On the other hand, African origins have been envisaged for the 1836; extinct only few centuries ago), the Seychelles Archipelago sole other described reptilian inhabitant of Scontrone, i.e., the cro- and Madagascar (Aldabrachelys Loveridge and Williams, 1957; codylid cf. Crocodylus, which has been suggested to represent the Quaternary, including also an extant taxon), the Caribbean Sea product of a direct or indirect marine dispersal from Africa that col- and the Bahamas Archipelago (Chelonoidis; Quaternary), and cer- onized the palaeoisland of Scontrone during the Tortonian (Delfino tain more poorly known taxa from the Canary Islands (‘‘Cen- and Rossi, 2013). trochelys”; Plio-Pleistocene), the Cape Verde Islands In any case, fossil record, molecular data, as well as direct obser- (‘‘Centrochelys”; Pleistocene), Indonesia (?Megalochelys Falconer vations of extant taxa, attest that Mauremys and, especially, large and Cautley, 1837; Pleistocene), and Philippines (?Megalochelys; testudinids are well capable of dispersing throughout large marine Pleistocene) (Günther, 1877; Van Denburgh, 1914; Arnold, 1979; distances (Gerlach et al., 2006; Vamberger et al., 2014; Kehlmaier Gerlach, 2004; Setiyabudi, 2009; Rhodin et al., 2015; Vlachos, 9 G.L. Georgalis and M. Delfino Geobios xxx (xxxx) xxx 2018; Kehlmaier et al., 2019; Georgalis et al., 2021). A principal Blain, H.-A., Fagoaga, A., Ruiz-Sánchez, F.J., Bisbal-Chinesta, J.F., Delfino, M., 2019. Latest Villafranchian climate and landscape reconstructions at Pirro Nord common feature of almost all of these endemic insular testudinid (southern Italy). Geology 47, 829–832. forms is the acquisition of a large size. These being said, the Scon- Chesi, F., Delfino, M., Abbazzi, L., Carboni, S., Lecca, L., Rook, L., 2007. New fossil trone testudinid could be a descendant of any of the late Paleogene vertebrate remains from San Giovanni di Sinis (Late Pleistocene, Sardinia): the or Neogene continental large forms that inhabited Europe and last Mauremys (Reptilia, Testudines) in the central Mediterranean. Rivista Italiana di Paleontologia e Stratigrafia 113, 287–297. Afro-Arabia, or even of some smaller continental form that eventu- Chesi, F., Delfino, M., Rook, L., 2009a. Late Miocene Mauremys (Testudines, ally evolved a large size during its insular isolation. More remains Geoemydidae) from Tuscany (Italy): evidence of terrapin persistence after a are definitely needed in order to decipher the exact taxonomic mammal turnover. Journal of Paleontology 83, 379–388. Chesi, F. 2009b. Il registro fossile italiano dei cheloni. Ph.D. Thesis, Università di affinities and biogeographic origins of the Scontrone turtles. Firenze, Firenze, Italy. 178 pp. We are still only at the beginning of comprehending these Claude, J., Tong, H., 2004. Early testudinoid turtles from Saint-Papoul, France, with unique insular herpetofaunas on these now long-lost islands of comments on the early evolution of modern Testudinoidea. Oryctos 5, 3–45. Claude, J., Tong, H., Suteethorn, V., 2007. Turtles from the late Eocene–early the Central Mediterranean. Even though poorly preserved, the Gar- Oligocene of the Krabi Basin (Thailand). 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Late Messinian mollusks and vertebrates from Moncucco through this part of the Central Mediterranean (Georgalis et al., Torinese, north-western Italy. Paleoecological and paleoclimatological 2020). implications. Palaeontologia Electronica 20.1.10A, 1–66. Delfino, M., 2002. Erpetofaune italiane del Neogene e del Quaternario. Ph.D. Thesis, University of Modena and Reggio Emilia, Modena and Reggio Emilia (unpubl.). Declaration of Competing Interest Delfino, M., Böhme, M., Rook, L., 2007. First European evidence for transcontinental dispersal of Crocodylus (late Neogene of southern Italy). Zoological Journal of the The authors declare that they have no known competing finan- Linnean Society 149, 293–307. Delfino, M., Bailon, S., 2000. Early Pleistocene herpetofauna from Cava Dell’ Erba and cial interests or personal relationships that could have appeared Cava Pirro (Apulia, Southern Italy). Herpetological Journal 10, 95–110. to influence the work reported in this paper. 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