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New fossil postcranial remains of Testudo (Chersine) catalaunica from Castell de Barber a (CB), Hostalets de Pierola Superior (els Hostalets de Pierola, HPS) and Can Mata indeterminate (CM). (A, B) proximal fragment of right femur IPS36396b from CB in dorsal (A) and ventral (B) views. (C, D) proximal fragment of left femur IPS30905 from CM in dorsal (C) and ventral (D) views. (E, F) proximal fragment of left femur IPS87219 from CM in dorsal (E) and ventral (F) views. (G, H) proximal fragment of left femur IPS16441 from HPS in dorsal (G) and ventral (H) views.
Source publication
We provide a taxonomic review of the extinct testudinid Testudo catalaunica, based on published and unpublished material from several Miocene (late Aragonian and early Vallesian) sites of the Vallès-Penedès Basin (north-east Iberian Peninsula). We show that Testudo catalaunica irregularis is a junior subjective synonym of T. catalaunica, and furthe...
Contexts in source publication
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... 2, the pygal, and all the posterior peripheral plates are missing from the carapace, whereas most of the proximal portions of the left and right costals 1-4 and of left peripherals 4-7 are very cracked and chipped. Only the anterior edge of the epiplastra and xiphiplastra are missing from the plastron. IPS30905: Proximal fragment of a left femur (Fig. 8C, D), 1.51 cm long and 0.83 cm wide. It preserves most the proximal portion of the shaft, the head, the greater and lesser trochanters, and the intertrochanteric fossa. The ventral surface of the preserved bone is partially covered by matrix. (Fig. 8E, F), 1.25 cm long and 1.33 cm wide. It only preserves the femoral head, the two ...
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... and xiphiplastra are missing from the plastron. IPS30905: Proximal fragment of a left femur (Fig. 8C, D), 1.51 cm long and 0.83 cm wide. It preserves most the proximal portion of the shaft, the head, the greater and lesser trochanters, and the intertrochanteric fossa. The ventral surface of the preserved bone is partially covered by matrix. (Fig. 8E, F), 1.25 cm long and 1.33 cm wide. It only preserves the femoral head, the two trochanters, and the intertrochanteric fossa, which is completely filled by ...
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... Partial left femur (Fig. 8G, H), 2.25 cm long and 1.73 cm wide, which preserves the femoral head, the two trochanters, the intertrochanteric fossa, and a larger portion of the shaft than the two preceding ...
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... plates cannot be confidently assessed. The right costals 2-5 are also clearly displaced, overlapping the neural series. In contrast, the remaining fractures do not considerably dislocate the rest of the plates. IPS36396b is the only associated postcranial element, consisting of a proximal fragment of right femur, 0.91 cm long and 0.63 cm wide (Fig. 8A, B); the distal portion of the shaft is missing, and the intertrochanteric fossa is partially covered by sediment. . Like in other species of Testudo, the carapace displays no sculptur- ing; merely, scute growth lines can be discerned in IPS36396a (Fig. 6G) thanks to its good state of preservation (this is unusual in fossil ...
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... preserved limb elements only consist of proximal fragments of right (IPS36396b; Fig. 8A, B) and left (IPS30905, IPS87219, and IPS16441; Fig. 8C-H) femora. IPS36396b was found associated with the almost complete shell IPS36396a from CB, and the remaining femoral specimens are also attributed to T. catalaunica not only based on size and shape con- gruence with IPS36396b, but also because this spe- cies is the only ...
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... preserved limb elements only consist of proximal fragments of right (IPS36396b; Fig. 8A, B) and left (IPS30905, IPS87219, and IPS16441; Fig. 8C-H) femora. IPS36396b was found associated with the almost complete shell IPS36396a from CB, and the remaining femoral specimens are also attributed to T. catalaunica not only based on size and shape con- gruence with IPS36396b, but also because this spe- cies is the only medium-sized testudinid recorded in the area of els Hostalets de ...
Citations
... Second, the taxonomy of fossil tortoises is inconsistent, in part because fossil taxa have a history of being grouped and named based on body size alone [63]. Although the taxonomy of fossil tortoises has been revised in great detail in some regions and illuminated phylogenetic relationships of fossil taxa [18,61,62,[64][65][66][67][68][69], there are still many fossil records that have not undergone recent taxonomic reevaluation. Such additional study is necessary to place the specimens into a broader phylogenetic context with confidence (for discussion of this issue see [18]). ...
The late Quaternary is characterized by the extinction of many terrestrial megafauna, which included tortoises (Family: Testudinidae). However, limited information is available on how extinction shaped the phenotype of surviving taxa. Here, based on a global dataset of straight carapace length, we investigate the temporal variation, spatial distribution and evolution of tortoise body size over the past 23 million years, thereby capturing the effects of Quaternary extinctions in this clade. We found a significant change in body size distribution characterized by a reduction of both mean body size and maximum body size of extant tortoises relative to fossil taxa. This reduction of body size occurred earlier in mainland (Early Pleistocene 2.588–0.781 Ma) than in island tortoises (Late Pleistocene/Holocene 0.126–0 Ma). Despite contrasting body size patterns between fossil and extant taxa on a spatial scale, tortoise body size showed limited variation over time until this decline. Body size is a fundamental functional trait determining many aspects of species ecologies, with large tortoises playing key roles as ecosystem engineers. As such, the transition from larger sized to smaller sized classes indicated by our findings likely resulted in the homogenization of tortoises' ecological functions and diminished the role of tortoises in structuring the vegetation community.
... Despite its incompleteness, MT-VII-must-T-01 can be referred to Testudinidae and in particular to Testudo hermanni on the basis of the following characters (Amiranashvili 2000;Hervet 2000;Delfino et al. 2012;Luján et al. 2016;Vlachos and Rabi 2018): trapezoidal costal elements with ends alternatively broad and narrow; overlap of the pleuromarginal sulci with the costoperipheral sutures; ventral gular surface flat; epiplastral lips not particularly high and not overhanging posteriorly (they do not overhang the entoplastron) so that they do not develop a significant gular pocket. ...
We review in detail the published fossil record of turtles from Sardinia and, in addition, we document previously undescribed specimens for the first time. Among these undescribed specimens, is the oldest occurrence of Testudo hermanni on the island, from the Early Pleistocene of Monte Tuttavista. The turtle fossil record in Sardinia goes back to the Eocene and comprises 18 different taxa, pertaining to 6 lineages: Podocnemididae, Cheloniidae, Emydidae, Geoemydidae, Testudinidae, and Trionychidae. Remarkable is the occurrence of Eocene pleurodires, whose presence is in agreement with the Oligo-Miocene rifting of the Corso-Sardinian block. Interestingly, the fossil record provides evidence for the presence in the island of both Testudo hermanni and Emys orbicularis during the Pleistocene although according to molecular data the extant populations of these two taxa were introduced in recent times. Finally, a large ungual phalanx from the Middle–Late Pleistocene Monte San Giovanni bone breccia testifies the occurrence of a giant tortoise in the Quaternary terrestrial ecosystem of Sardinia.
... Chelonian fossils include scarce remains of the geoemydid Ptychogaster as well as a few disarticulated shell plates and some phalanges attributed to the medium-sized tortoise Testudo. The lack of diagnostic shell parts precludes an assignment to the subgenus rank for both taxa (Luján et al. 2014a(Luján et al. , 2016. The most abundant and complete chelonian material, comprising both shell and postcranial remains, belongs to giant tortoises of the genus Titanochelon. ...
Els Casots is one of the richest fossil vertebrate sites of the Vallès-Penedès Basin (Catalonia, Spain). It was
discovered in 1989 and excavated briefly during the 1990s, resulting in the recovery of thousands of remains
and the erection of several new mammal species. Excavations were resumed in 2018 and continue to date.
Here we provide updated results regarding the age, stratigraphy, biota and palaeoenvironment of the site.
The age of the site is well constrained to ~15.9 Ma thanks to recent bio- and magnetostratigraphic data, thus
coinciding with the Miocene Climatic Optimum (MCO). The stratigraphic succession at the site area indicates
lacustrine to palustrine environments with cyclically oscillating water level. There are several fossiliferous
layers that have yielded a vertebrate fauna comprising up to 75 different vertebrate species including
amphibians, reptiles, birds and mostly mammals. The finding of several articulated partial skeletons indicate
that the site records an autochthonous to parautochthonous assemblage. The abundance and completeness
of the vertebrate remains together with a well-constrained age and detailed stratigraphic and palaeoenvironmental
data, make els Casots a key site for understanding wetland ecosystems in southern Europe during
the MCO.
... On the other hand, the diverse testudinid records from Greece also bring important information on the phylogenetic relationships of the family Testudinidae. Although the taxonomy of many extinct members of this family remained problematic, the renewed interest on tortoise fossils from Europe and Greece over the last years has offered plenty of opportunities to investigate further the phylogeny of the tortoises (Pérez-García and Vlachos 2014;Vlachos 2015;Vlachos and Tsoukala 2016;Luján et al. 2016;Pérez-García et al. 2017a;Vlachos and Rabi 2018). Now it becomes clear that derived tortoises from Europe and Greece (i.e., Testudininae) belong mainly to two clades: Geochelonini and Testudinini. ...
... Included Taxa Extant members of Testudo include the Testudo graeca complex (with at least five subspecies), Testudo marginata, and Testudo kleinmanni. Several extinct species have been named or considered as members of Testudo, but few of them can actually be demonstrated to be members of this clade (see Luján et al. 2016;Vlachos and Rabi 2018). It is however clear that this clade has a fossil record that extends at least till the Late Miocene, as the Late Miocene Testudo marmorum and Testudo hellenica from Greece are certainly the oldest members of this clade. ...
... Other extant small testudinids that have been traditionally under Testudo (hermanni, horsfieldii) are considered as part of distinct genera, Chersine and Agrionemys, respectively. These genera are considered as subgenera of Testudo by Luján et al. (2016). ...
Turtles are among the most common and easily identifiable elements of the vertebrate fossil record, especially in continental deposits. In Greece, turtle fossils have been identified in at least 68 continental and insular localities, ranging from the Early Miocene to Holocene deposits. The review of the known turtle fossil record from Greece reveals the presence of at least 14 different species belonging to 4 cryptodiran clades (Trionychidae, Emydidae, Geoemydidae, and Testudinidae) and a single pleurodiran clade. The most diverse clade is Testudinidae, comprising more than half of the known diversity, with at least two giant and seven small-sized tortoise species. Although turtle fossils from Greece have been known since the first collections in the middle of the nineteenth century, they have received special attention only recently.
... Both of these clades are documented in the Afro-Arabian fossil record, with the majority belonging to Geochelona (Figure 1). Several Afro-Arabian fossil testudinid taxa have been placed into phylogenetic analyses, such as Gigantochersina ammon, Impregnochelys pachytectis, Namibchersus namaquensis, Mesochersus orangeus, Testudo kenitrensis, and Testudo oughlamensis (Gaffney and Meylan 1988;Gmira 1995;Luján et al. 2016;Vlachos and Rabi 2018;Pérez-García et al. 2020;), but relationships often remain far from resolved. We refrain from assigning "Geochelone" laetoliensis and "Geochelone" stromeri to either geochelonans or testudonans (although they probably pertain to the former clade), and, instead for the sake of convenience, we refer them to the "wastebasket" genus Geochelone Fitzinger, 1836, although this generic placement is certainly not correct. ...
... Delfino, unpublished data). The lineage of the extant species Testudo kleinmanni Lortet, 1883, can be dated back to the Pliocene, as a recent phylogenetic analysis placed the extinct T. kenitrensis and T. oughlamensis as its sister taxa (Luján et al. 2016). Furthermore, on the basis of molecular data, Testudo graeca, which is currently found on both sides of the Mediterranean, has been suggested to have had an Eastern European origin and then dispersed to northern Africa during the Pliocene, while its current population in the Iberian Peninsula and Italy are the product of a much later dispersal during historic times ). ...
... Comments. The genus Testudo has recently been partitioned into different lineages, which, depending on taxonomic preferences, have been either treated as subgenera or independent genera (Lapparent de Broin et al. 2006aBroin et al. , 2006bParham et al. 2006;Luján et al. 2016;TTWG, 2017;Vlachos and Rabi 2018;Garcia et al. 2020). We here follow the taxonomic scheme of TTWG (2017), in which Agrionemys Khosatzky and Młynarski, 1966 and Chersina Merrem, 1820 are treated as subgenera of Testudo. ...
Turtles of the clade Testudinoidea have a rather scarce fossil record in Afro-Arabia, ranging from the late Eocene up to the Quaternary. The vast majority of testudinoid fossils from Afro-Arabia are ascribed to Testudinidae, which has had a continuous presence in the area since the late Eocene. Geoemydidae is poorly documented by fragments found throughout the Neogene across northern Africa and the Middle East. Emydidae is absent from the fossil record of this area. All valid named taxa pertain to testudinids. Within Testudinidae, the majority of known fossil species are members of the clade Geochelona, while a few others belong to the clade Testudona. Four fossil taxa are members of now-extinct genera, five are members of extant genera, and seven cannot be assigned to a known genus with certainty. The fossil record also documents that several extant genera had a much broader distribution during the Neogene and Quaternary. Endemic insular lineages were formerly present on the Canary Islands, Cape Verde islands, and on several islands in the Western Indian Ocean. The highest known diversity of testudinoids seems to have existed during the Neogene; however, definitive conclusions are hampered by the extremely poor Paleogene record and large, unsampled areas of Afro-Arabia. A taxonomic review of the 22 named Afro-Arabian taxa finds 16 nomina valida, 1 nomen invalidum, and 5 nomina dubia.
... Paleotestudo canetotiana (Lartet, 1851) from the middle Miocene of Sansan (France) and Testudo antiqua Bronn, 1831 from the middle Miocene Hohenhöwen (Germany) were considered as representatives of the same 'hermanni lineage' (Lapparent de Broin et al. 2006a, p. 708). Subsequent phylogenetic studies confirmed the basal position of Paleotestudo canetotiana in the hermanni clade (Corsini et al. 2014;Pérez-Garcia et al. 2015;Luján et al. 2016), whereas the position of Testudo antiqua varied between polytomy with the extant clades of Testudo sensu lato (Corsini et al. 2014) to a variable position in the hermanni clade (Pérez- Luján et al. 2016). The attribution of T. antiqua to the genus Paleotestudo is confirmed by Pérez-García (2016). ...
... Paleotestudo canetotiana (Lartet, 1851) from the middle Miocene of Sansan (France) and Testudo antiqua Bronn, 1831 from the middle Miocene Hohenhöwen (Germany) were considered as representatives of the same 'hermanni lineage' (Lapparent de Broin et al. 2006a, p. 708). Subsequent phylogenetic studies confirmed the basal position of Paleotestudo canetotiana in the hermanni clade (Corsini et al. 2014;Pérez-Garcia et al. 2015;Luján et al. 2016), whereas the position of Testudo antiqua varied between polytomy with the extant clades of Testudo sensu lato (Corsini et al. 2014) to a variable position in the hermanni clade (Pérez- Luján et al. 2016). The attribution of T. antiqua to the genus Paleotestudo is confirmed by Pérez-García (2016). ...
... The recent revision of Ch. lunellensis by Delfino et al. (2012) resulted in the first diagnosis and provided additional material of this species, that supported its systematic position. Luján et al. (2016) additionally suggest to place into Chersine (as Testudo (Chersine)) the following Miocene species: Testudo escheri Pictet et Humbert, 1856 from the middle Miocene of Winterthur (Switzerland); T. mellingi (Peters, 1868) Młynarski, 1983 from the late Miocene of du Kohfidisch (Austria); T. opisthoklitea Schleich, 1981 from the middle Miocene of Gammelsdorf (Germany); and T. rectogularis Schleich, 1981 from the transitional early/middle Miocene of Sandelzhausen (Germany). ...
The shell remains of tortoises of the genus Chersine (= Eurotestudo) are described from the early Pliocene of the Musait and Colibasi localities of Moldova. The incomplete shell from the Musait locality is assigned to a new species, Chersine khosatzkyi sp. nov., which can be distinguished from all other species of Chersine by relatively wide posterior lobe of the plastron with nearly parallel lateral borders and presence of additional suprapygal. An isolated pygal from the Colibasi locality is attributed to Chersine sp. Chersine from the Musait and Colibasi localities represent first reliable records of Chersine in Moldova and the easternmost records of Chersine (the second one outside the Mediterranean region). In addition, these finds are among the oldest fossil records of the genus.
... In addition, the large, shallow, triangular depression above the condyles, pierced by a small foramen that opens on the proximal rim of the depression are characteristic of varanid lizards. According to Lécuru (1969) and Smith (2009) this foramen is observed in nearly all limbed squamates, but it is quite reduced or even absent in Varanus. Georgalis et al. (2018) described an humerus from the late Miocene of Ravin de la Pluie (Greece). ...
Amphibians and reptiles are decribed for the first time from the Cenozoic of Afghanistan. They originate from four Neogene localities. Sherullah 9 (late Miocene) yielded anuran amphibians (? Alytidae Fitzinger, 1843 (? Discoglossinae Günther, 1858), “Ranidae” Batsch, 1796 and another, indeterminate family), one chelonian, the terrestrial testudinine Agrionemys Khozatsky & Młynarski, 1966, which is compared to a Maragheh fossil specimen, and the genus still being present in the Kabul area; one indeterminate lizard, snakes (Colubridae Oppel, 1811 s.l. and two distinct snakes that pertain to either the Colubridae s.l. or Elapidae Boie, 1827). The faunas of the three other localities are very
poor. Molayan (late Miocene) produced only one lizard (Varanus sp. Merrem, 1820, Varanidae Gray, 1827). Pul-E Charkhi (early Pliocene) yielded indeterminate anurans, one indetermined lizard and colubrid snakes. Only an indeterminate anuran family is known at Hadji Rona (early Pliocene). Then, as known, the fauna is constituted of families which still have representatives living in Afghanistan.
... Both AMPG 1999 and GNHM 22 are morphologically consistent with members of the Testudona clade. Unfortunately, however, the diagnostic presence/absence of a hypo-xiphiplastral hinge (see Lapparent de Broin et al., 2006;Luján et al., 2016;Vlachos and Tsoukala, 2016;Vlachos and Rabi, 2018) cannot be confirmed in either specimen. Nevertheless, AMPG 1999 exhibits a characteristically symmetrical suture between costal 7 and suprapygal 1; this encloses costal 8 and excludes it from contact with neural 8 (Fig. 2D). ...
The world-famous upper Miocene fossil localities on the Aegean island of Samos in Greece have produced a rich fossil record that sheds light on the evolution of eastern Mediterranean terrestrial faunas over a one-million-year interval of the late Neogene. Fossils have been discovered on Samos since antiquity, although a succession of paleontological and commercial collecting expeditions over the last 130 years has resulted in specimens now being distributed throughout museums all over the world. Here, we survey the fossil tortoise remains from Samos, which are significant because they include early antecedents of the modern Testudo lineage, together with spectacular examples of the European Neogene gigantic testudinid †Titanochelon, which represents one of the largest-bodied terrestrial turtle taxa documented to date. All of the Samos fossils derive from the Mytilinii Formation, which spans the late MN11–early MN13 Neogene land mammal zones. The small-bodied tortoise remains include two incomplete shells that are morphologically consistent with basal testudonans and phylogenetically distinct from the coeval species Testudo marmorum found on mainland Greece. The Samos gigantic tortoise †‘Testudo’ schafferi was based on a spectacularly large skull and femur. However, we describe new plastron fragments, limb elements, and osteoderms that are compatible with †Titanochelon specimens from southern Greece and Anatolia. This could imply faunal links with the distinctive ‘Pikermian’ local assemblages from Asia Minor and concurs with the proposed late Miocene–Pliocene biogeographic segregation of large mammals from the eastern Aegean margin and Turkey relative to those occurring in northwestern Greece and the Balkan Peninsula.
... In order to observe the detail of the anatomical elements, the specimen was completely prepared. In order to strengthen the systematic attribution of the fossil and discuss its affinities, a phylogenetic analysis was performed based on a published morphological matrix (Table 1), which includes 20 taxa and 37 characters [32] that helped to resolve the relationships between the various Testudo s. l. based on morphological characters. The definition and the coding of some characters were modified (S1 Table). ...
... Table 1. Data matrix of the cladistic analysis, modified after Lujan et al [32]. Missing data are represented by a question mark. ...
We here report on fossil remains of the earliest known crown-Testudo, an extant clade of Mediterranean testudinid tortoises from the late Miocene (Vallesian, MN 10) from the hominoid locality Ravin de la Pluie (RPl) in Greece. The material studied is a small, nearly complete carapace with a clearly distinct hypo-xiphiplastral hinge. This supports the sensu stricto generic assignment. This new terrestrial testudinid specimen is characterized by a possible tectiform, narrow, elongated shell with a pentagonal pygal and a long, posteriorly elevated, lenticular and rounded dorsal epiplastral lip. These unique features differ from those of other known Mediterranean hinged forms and allow the erection of the new species Testudo hellenica sp. nov. This taxon is phylogenetically close to two Greek species, the extant T. marginata and the fossil T. marmorum (Turolian, around 7.3 Ma). This record provides evidence for the first appearance of the genus Testudo sensu stricto at a minimum age of 9 Ma.
... Phylogenetic Position-Recently, the phylogenetic position of Testudo marmorum was analyzed in several publications Luján et al., 2016;Vlachos and Tsoukala, 2016;Vlachos and Rabi, 2018). All analyses agree that Te. marmorum holds a derived position within Testudo and that it belongs to the subclade Chersus (in Parham et al., 2006a), together with Testudo marginata and Testudo kleinmanni. ...
... All analyses agree that Te. marmorum holds a derived position within Testudo and that it belongs to the subclade Chersus (in Parham et al., 2006a), together with Testudo marginata and Testudo kleinmanni. Whereas Corsini et al. (2014) recover it as the sister taxon of Te. marginata and Luján et al. (2016) recover it as sister to the Te. marginata + Te. brevitesta clade (both are analyses of morphological characters), Vlachos and Tsoukala (2016; morphological characters with constrained molecular topology) and Vlachos and Rabi (2018;total-evidence analysis) suggest that it is the sister taxon of Te. kleinmanni. ...
... Another alternative is proposed by Parham et al. (2006a), who explain the distribution of the mentioned species on either side as a result of vicariance after the separation of the two areas in post-Messinian times. Finally, the topology retrieved by Luján et al. (2016) provides a logical, from a chronostratigraphic point of view, position of Te. brevitesta as well. ...
This article deals with the fossil tortoises of one of the most iconic fossil localities of the Neogene of the Old World, the upper Miocene locality of Pikermi, near Athens, Greece. We describe the type, previously published, and new material of the fossil tortoises from Pikermi, along with new material from the coeval Azmaka 6 locality in Bulgaria. This combined information results in the complete revision of the ‘marble tortoise’ Testudo marmorum, which is the first turtle species ever named from Greece. Besides the challenges and difficulties of working with material from the old collections of Pikermi, we present a complete revision of this species and identify its presence outside its type locality in the South Balkan Peninsula. The presence of numerous shells of the ‘marble tortoise’ allows us to discuss the observed variation and attempt to interpret it as the result of intraspecific variation, ontogenetic changes, or sexual dimorphism. For the first time, we observe the absence of the characteristic hypo-xiphiplastral hinge in female individuals of Testudo marmorum, which in turn challenges the traditional phylogenetic position of this species. We further describe previously published and new material of the giant tortoise Titanochelon from Pikermi. Based on the updated information, we can suggest that the Pikermian giant tortoise probably represents a distinct taxon. Our results signal Pikermi as one of the most important localities to understand the evolution of tortoises in the eastern Mediterranean during the last parts of the Neogene.
