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SOLITUDO SICULA SP. NOV. FIGS 4–5 Zoobank registration: urn: lsid: zoobank. org:act: F5011B57-E559-4F72-9131-D457D8B6B998. Holotype: ZCSP US.0-Q2, an almost complete right femur, missing only part of the major trochanter (Fig. 4). Paratypes: ZCSP US.0-Q1, a fragment of a right ischium (Fig. 5A, B); ZCSP US.2-Q4, a left pubis (Fig. 5C, D); ZCSP US.0-Q3, an ungual phalanx (Fig. 5 E-G). Due to the uncertainty of the attribution of the four skeletal remains to a single individual (see below), we do not group the ischium, the pubis and the phalanx along with the femur as a single holotype, but we keep them separated. Type locality and age: Zubbio di Cozzo San Pietro Cave, Municipality of Bagheria, near Palermo, northwestern Sicily, Italy. Latest Late Pleistocene. For details about the locality, see Locality information above. The tortoise remains here described come from an archaeological funerary area (Battaglia et al., 2020) placed at a depth of − 15.4 m from the entrance and characterized by disturbance caused by human and/or animal activity that determined a complex stratigraphy. The remains of the pubis, ZCSP US.2-Q4, was AMS-dated revealing an age of 12.5 ± 0.5 kya BP (Supporting Information, Fig. S3). All the tortoise material is referred to the latest Late Pleistocene assuming that it belongs to the same, extremely rare taxon (large-sized tortoises have never been reported in the several Sicilian Late Pleistocene-Holocene archaeological and paleontological sites so far studied), even if not necessarily to a single specimen (considering the alteration of the stratigraphy, a non-demonstrable but likely option because of non-anatomical overlap of the preserved skeletal elements, as well as their congruent size and common testudinid morphology). Etymology: Latin adjective in feminine gender, meaning ‘Sicilian’. Diagnosis: Member of Solitudo based on the generic characters mentioned above. Differing from other species of Solitudo in having trochanters that are at the same height with the femoral head, and a femoral head that is clearly elliptical, whereas in other species trochanters extend beyond the femoral head and the latter is not elliptical. Description Femur: ZCSP US.0-Q2 (Fig. 4; Supporting Information, Fig. S4) is a well-preserved right femur with a total length of 104 mm. The diaphysis is relatively slender and almost rectilinear, with a slight curvature on the distal part [more rectilinear than femora of Te. hermanni (adult and young individuals) and Te. marginata]. The dimensions of the narrowest point of the diaphysis are 13.1 × 12.9 mm. It is not possible to measure the original width of the femur at the level of the trochanters, as the trochanter major is laterally incomplete. The width of its preserved portion is 32 mm. The maximum width of the distal epiphysis is 37 mm. The neck is well defined, relatively long and contracted with respect to the maximum width of the head (not as in the Menorcan species which has a femoral head detached even more clearly from the neck). The minimum width of the neck is 22 mm; its minimum thickness is 16 mm. The intertrochanteric fossa is wide and occupies the entire surface between the trochanters; it is approximately 10 mm deep. The trochanters do not extend more proximally compared to the femoral head (see Supporting Information, Appendix S1 for comments on the orientation of the femur). The lateral surface of the head neck is nearly vertical, the head does not overhang the shaft as in Te. hermanni and Te. marginata. The two trochanters are joined ventrally (so that the intertrochanteric fossa does not open ventrally), but the muscular insertion surfaces are clearly separated, and in medial view, the space between these two surfaces becomes thinner until it forms a relatively thin crest. In ventral view, this crest is not rectilinear but forms a wide ‘U’ (above which much of the head of the femur is visible). These areas of muscular insertion are clear, as they are defined by a thin ridge. A similar ridge delimits the ventral margin of the muscular insertion surface of the head—the one that overlooks the intertrochanteric fossa; another similar crest delimits the insertion region of the distal condyles. The area of muscular insertion of the small trochanter is clearly triangular, and probably also that of the great trochanter, although the latter is partially incomplete. On the anterior surface of the diaphysis of the femur, at the small trochanter, there is a long ridge of muscular insertion of the femorotibialis (more than half of the diaphysis) evident but moderately developed (although not as much as in Te. marginata); it remains close to the edge between the anterior and ventral surface. On the posterior surface of the diaphysis, in connection with the great trochanter, there is an elongated area (depression with an irregular base) of muscular insertion of the adductor femoris, which expands distally by curving in a dorsal direction until it reaches the dorsal edge of the posterior surface at about the half of the diaphysis. The femoral head is narrow, narrower than the combined width of the trochanters (even more if we account for the missing part) and oval, clearly elliptical in outline, being oriented at an angle of approximately 30° compared to the anteroposterior plane. Distally, the tibial and fibular condyles are connected by a ridge ventrally, which is gently curved but only weakly developed. Ischium: ZCSP US.0-Q1 (Fig. 5A, B; Supporting Information, Fig. S5) is an incomplete right ischium, missing the anteromedial area that contacts with the contralateral element and with the pubis. The anterior margin is laminar (it forms a sharp crest). On its ventral surface, at this anterior laminar margin there is a moderate depression medially bounded by an evident termination. The posterior margin is rounded and in dorsal view it forms a wide concavity. The dorsal surface near the medial margin is slightly concave. Posteromedially, the lateral ischial process forms a sort of low and broad tubercle with a rough surface (note that in Te. hermanni and Te. marginata, the morphology is completely different, taking the shape of a pointed process). Near the posteromedial margin, the ischium thickens ventrally as it is also indicated by the shape of the suture area with the left ischium. The posterior surface of this thickened area is characterized by an area of muscular insertion delimited by a small ridge similar to that of the trochanters of the femur. Pubis: ZCSP US.2-Q4 (Fig. 5C, D) was an incomplete right pubis, preserving only its posterolateral portion. The three articular facets and the laminar region leading to the two other processes are preserved. This element has been destructively sampled for AMSdating and no longer exists. U n g u a l p h a l a n x: Z C S P U S.0- Q 3 (F i g. 5E - G; Supporting Information, Fig. S6) is an ungual phalanx, with a length of 29 mm. The articular facet is divided into two concavities separated by a vertical convexity. The upper portion of the proximal epiphysis is clearly projecting. On its ventral surface, near the facet joint, there are two prominent foramina. The gross morphology of the phalanx is clearly different from that of Testudo spp.; it is wide and flat instead of being rounded in section. Size: The femur is much larger than that of Te. marginata, which is the largest extant tortoise species in Europe (Ernst & Barbour, 1989; Bringsøe et al., 2001). For example, the Te. marginata femur MDHC 370 has a length of 51 mm, while the straight carapace length of the same specimen is 300 mm. On the basis of the length difference of the femora (ZCSP US.0-Q2 is 104 mm long), we estimate the straight carapace length of the extinct Sicilian tortoise as exceeding 50 cm and maybe up to 60 cm, i.e. significantly larger than that of extant Te. hermanni [up to 37.5 cm in the Balkans and up to 22 cm in the western Mediterranean (Cheylan, 2001)], or any other Testudo species. The size record for Te. marginata, the largest species of the genus, is 40 cm, but most specimens are significantly smaller (Bringsøe et al., 2001)., Published as part of Valenti, Pietro, Vlachos, Evangelos, Kehlmaier, Christian, Fritz, Uwe, Georgalis, Georgios L., Luján, Àngel Hernández, Miccichè, Roberto, Sineo, Luca & Delfino, Massimo, 2022, The last of the large-sized tortoises of the Mediterranean islands, pp. 1704-1717 in Zoological Journal of the Linnean Society 196 (4) on pages 1711-1714, DOI: 10.1093/zoolinnean/zlac044, http://zenodo.org/record/7390583, {"references":["Battaglia G, Micciche RM, Forgia V, Mannino MA, Scopelliti MG, Sineo L, Valenti P. 2020. New perspectives on the human occupation of the Gulf of Palermo during the Metal Ages: the funerary cave of Zubbio di Cozzo San Pietro (Bagheria) and the necropolis of Viale Venere (Mondello, Palermo). Journal of Archaeological Science: Reports 31: 102276.","Ernst CH, Barbour RW. 1989. Turtles of the world. Washington, DC: Smithsonian Institution Press.","Bringsoe H, Buskirk JR, Willemsen RE. 2001. Testudo marginata Schoepff, 1792 - Breitrandschildkrote. In: Fritz U, ed. Handbuch der Reptilien und Amphibien Europas, Band 3 / IIIA, Schildkroten (Testudines I) Bataguridae, Testudinidae, Emydidae. Wiebelsheim: Aula-Verlag, 291 - 334.","Cheylan M. 2001. Testudo hermanni Gmelin, 1789 - Griechische Landschildkrote. In: Fritz U, ed. Handbuch der Reptilien und Amphibien Europas, Band 3 / IIIA, Schildkroten (Testudines I) Bataguridae, Testudinidae, Emydidae. Wiebelsheim: Aula-Verlag, 179 - 289."]}

SOLITUDO GEN. NOV. Zoobank registration: urn: lsid: zoobank. org:act: 08DE006C-E90B-4605-9711-952D72904E 5F. Type species: Testudo robusta Leith-Adams, 1877; Zubbo Cave = Zebbug Cave, Malta; Middle Pleistocene. Etymology: From the Latin word solitudo, feminine, third declension, meaning solitude, loneliness, in allusion to the insular isolation of these tortoises. Also, the termination - tudo alludes to the name Testudo and the testudinid affinities of this new taxon. Diagnosis: Members of Solitudo belong to Testudinidae because of the ventral fusion of the trochanters of the femur. However, Solitudo species show an incomplete fusion of the trochanters dorsally, in contrast to other testudinid genera in which the trochanters are connected proximally via a rounded ridge. Furthermore, members of Solitudo have femurs that are slender, with a femoral head that is narrower than the combined anteroposterior width of the trochanters and an oblique orientation compared to the anteroposterior plane between 25°–45°, in contrast to other genera with relatively massive femora and broad and less oblique femoral heads, forming an angle that is less than 25° with the anteroposterior plane. Included taxa: Solitudo robusta (Leith-Adams, 1877) comb. nov., Solitudo gymnesica (Bate, 1914) comb. nov., Solitudo sicula sp. nov. Remarks: The exact placement of Solitudo among Testudininae is unclear and should be examined when new material (skull, shell)is discovered.The femur alone, in its distinctiveness, does not allow firm assessment of the phylogenetic relationship with respect to any potentially related taxon (Centrochelys, Stigmochelys, Testudo and Titanochelon). It is noteworthy that the establishment of new fossil taxa on the basis of appendicular elements is not rare among testudinids, and has been used, besides most other insular taxa mentioned already, for the recently described species Chelonoidis marcanoi Turvey et al., 2017 from the Quaternary of Hispaniola (Turvey et al., 2017)., Published as part of Valenti, Pietro, Vlachos, Evangelos, Kehlmaier, Christian, Fritz, Uwe, Georgalis, Georgios L., Luján, Àngel Hernández, Miccichè, Roberto, Sineo, Luca & Delfino, Massimo, 2022, The last of the large-sized tortoises of the Mediterranean islands, pp. 1704-1717 in Zoological Journal of the Linnean Society 196 (4) on page 1711, DOI: 10.1093/zoolinnean/zlac044, http://zenodo.org/record/7390583, {"references":["Leith-Adams A. 1877. On gigantic land-tortoises and a small freshwater species from the ossiferous caverns of Malta, together with a list of their fossil fauna; and a note on chelonian remains from the rock cavities of Gibraltar. Quarterly Journal of the Geological Society of London 33: 177 - 191.","Bate DMA. 1914. On remains of a gigantic land tortoise (Testudo gymnesicus n. sp.) from the Pleistocene of Menorca. Geological Magazine 1: 100 - 107.","Turvey ST, Almonte J, Hansford J, Scofield RP, Brocca JL, Chapman SD. 2017. A new species of extinct Late Quaternary giant tortoise from Hispaniola. Zootaxa 4277: 1 - 16."]}