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... In this analysis Ba. trani had a different position by being placed as sister-taxon to crown Geoemydidae. [Staesche, 1928]) and Geiseltal (Borkenia germanica [Hummel, 1935]), both in Germany, but their taxonomic status and variation are in need of revision. The second lineage ("Ptychogasteridae group" of Hervet, 2004b) is best represented by Geiselemys ptychogastroides (Hummel, 1935), also from Geiseltal, Germany. ...
... [Staesche, 1928]) and Geiseltal (Borkenia germanica [Hummel, 1935]), both in Germany, but their taxonomic status and variation are in need of revision. The second lineage ("Ptychogasteridae group" of Hervet, 2004b) is best represented by Geiselemys ptychogastroides (Hummel, 1935), also from Geiseltal, Germany. We mostly compare Banhxeochelys trani to these species, as they appear to be representative for the diversity of European geoemydids from the Eocene/Oligocene. ...
... This is generally true for fossils as well, but particularly fossil rich localities or formations often yield large samples of turtles that can be used to document skeletal variation in extinct species. Large numbers of fossil geoemydids (i.e., more than 10 individuals) have previously been reported from the Eocene of Messel, Germany (Cadena, Joyce & Smith, 2018), the Eocene of Geiseltal, Germany (Hummel, 1935), and the Eocene of Wyoming (Gilmore, 1945;Brand et al., 2000), but no study has of yet properly summarized and discussed intraspecific variation based on this material. ...
Geoemydidae is a major group of turtles with 71 living species and a rich fossil record from the Tertiary period of the Northern Hemisphere. Despite its great species richness, little is known about the evolutionary history of this group and the phylogenetic relationship among its species, particularly fossils. This is in part due to the high levels of variation (polymorphism) observed in geoemydids species, and in part due to the lack of species-exclusive characteristics (synapomorphies), that precludes the relationship of species to be established with confidence. In the past, highly polymorphic species were often omitted from phylogenetic studies, however in this study I reconstruct the phylogenetic relationship of fossil and living geoemydids species by embracing the use of polymorphism, and using only characters of the turtle shell, which is more frequently preserved in the fossil record. In addition, I describe a new species of fossil turtle that lived around 34 million years ago based on rich material from Vietnam, and estimate the evolutionary position of this species and other fossil geoemydids from the Paleocene, Eocene and Oligocene epochs of North America, Europe and Asia. At last, I review the species identification and taxonomic status of many fossil geoemydid specimens from the Neogene and Quaternary periods of India and Pakistan, as well as illustrate and briefly describe all these specimens that were poorly described in the late 19th and early 20th centuries.
... We here agree with Hervet (2004aHervet ( , 2004b and that two lineages are present in the Eocene, although we side with by seeing less taxonomic diversity. The first lineage ("Palaeochelys sensu lato-Mauremys" of Hervet, 2004a; Palaeochelys of is best represented by rich material from Messel (Palaeoemys messeliana; Staesche, 1928) and Geiseltal (Borkenia germanica; Hummel, 1935), both in Germany, but their taxonomic status and variation are in need of revision. The second lineage ("Ptychogasteridae group" of Hervet, 2004b) is best represented by Geiselemys ptychogastroides (Hummel, 1935), also from Geiseltal, Germany. ...
... The first lineage ("Palaeochelys sensu lato-Mauremys" of Hervet, 2004a; Palaeochelys of is best represented by rich material from Messel (Palaeoemys messeliana; Staesche, 1928) and Geiseltal (Borkenia germanica; Hummel, 1935), both in Germany, but their taxonomic status and variation are in need of revision. The second lineage ("Ptychogasteridae group" of Hervet, 2004b) is best represented by Geiselemys ptychogastroides (Hummel, 1935), also from Geiseltal, Germany. We mostly compare Banhxeochelys trani to these species, as they appear to be representative for the diversity of European geoemydids from the Eocene/Oligocene. ...
... This is generally true for fossils as well, but particularly fossil rich localities or formations often yield large samples of turtles that can be used to document skeletal variation in extinct species. Large numbers of fossil geoemydids (i.e., more than 10 individuals) have previously been reported from the Eocene of Messel, Germany (Cadena, Joyce & Smith, 2018), the Eocene of Geiseltal, Germany (Hummel, 1935), and the Eocene of Wyoming (Gilmore, 1945;Brand et al., 2000), but no study has of yet properly summarized and discussed intraspecific variation based on this material. ...
Background
Testudinoidea is a major clade of turtles that has colonized different ecological environments across the globe throughout the Tertiary. Aquatic testudinoids have a particularly rich fossil record in the Tertiary of the northern hemisphere, but little is known about the evolutionary history of the group, as the phylogenetic relationships of most fossils have not been established with confidence, in part due to high levels of homoplasy and polymorphism.
Methods
We here focus on describing a sample of 30 testudinoid shells, belonging to a single population that was collected from lake sediments from the middle to late Eocene (35–39 Ma) Na Duong Formation in Vietnam. The phylogenetic placement of this new material is investigated by integrating it and 11 other species of putative geoemydids from the Eocene and Oligocene to a recently published matrix of geoemydid turtles, that embraces the use of polymorphic characters, and then running a total-evidence analysis.
Results
The new material is highly polymorphic, but can be inferred with confidence to be a new taxon, Banhxeochelys trani gen. et sp. nov. It shares morphological similarities with other southeastern Asian testudinoids, Isometremys lacuna and Guangdongemys pingi , but is placed phylogenetically at the base of Pan-Testuguria when fossils are included in the analysis, or as a stem geoemydid when other fossils are deactivated from the matrix. The vast majority of other putative fossil geoemydids are placed at the base of Pan-Testuguria as well.
Discussion
The phylogenetic placement of fossil testudinoids used in the analysis is discussed individually and each species compared to Banhxeochelys trani gen. et sp. nov. The high levels of polymorphism observed in the new taxon is discussed in terms of ontogenetic and random variability. This is the first time that a large sample of fossil testudinoids has its morphological variation described in detail.
... The European testudinids 'Hadrianus' eocaenica (Hummel, 1935), from the Lutetian of Germany, and 'Hadrianus' sp. (sensu Jim enez Fuentes et al., 1988;Figure 1. Geographical position of the type localities of the European Eocene testudinids discussed here. ...
... gigas, proposed by P erez-Garc ıa (2015), is taken into account. 'Hadrianus' eocaenica is coded on the basis of information obtained from the specimens presented by Hummel (1935), including the holotype. The taxon from Mazater on is studied here for the first time, based on abundant material, which has been prepared for the present study . ...
... For character 8 (shape of the third neural), two new states are included: hexagonal, short sides positioned anteriorly (state 3; Figs 2A, 3B, C, D); hexagonal, short sides positioned posteriorly (state 4; fig. 23 in Hummel, 1935). For character 11 (contact between the suprapygals), a new state is included: fused suprapygals (state 2). ...
Testudinidae is a clade of tortoises known in Europe from the early Eocene. It is the only lineage of terrestrial turtles identified in the post-Paleocene record of this continent. However, very little information on the taxonomy and systematics of the Eocene representatives is currently available. The oldest testudinids from Europe, recovered in several countries, are studied here. Some of them were previously attributed to genera defined in North America. The hypothesis suggesting that the European Eocene testudinids correspond to genera exclusive to this continent is confirmed here. In this sense, two new genera, Fontainechelon (early Eocene) and Pelorochelon (middle Eocene), are proposed. The largest collection of European Paleogene testudinids, from the middle Eocene of the Spanish site of Mazaterón (Soria), is analyzed for the first time. A new species from this locality is defined. The validity of various previously proposed taxa is refuted, and the diagnosis of several forms is emended. Thus, the diversity, taxonomy, phylogenetic relationships, and biogeographical and stratigraphic distributions of the European primitive representatives of this successful extant clade are analyzed.
... 1. Neurals with short postero-lateral sides appeared first in Early to Middle Eocene in both Geoemydidae (Geoemyda saxonica and Geoemyda ptychogasteroides (Hummel 1935)) and Testudinidae (Hadrianus, Achilemys). ...
... First species of this group may be "Geoemyda" ptychogasteroides from the middle Eocene of Germany (Hummel 1935) or "Ocadia" parisiensis from the middle Eocene of France (Botez 1921). ...
... Non exclusive synapomorphies within Testudinoidea: characters 40 and 42. The oldest known species belonging to this group may be "Geoemyda" ptychogasteroides from the Middle Eocene of Germany (Hummel 1935) or "Ocadia" parisiensis from the Middle Eocene of France (Botez 1921). ...
... Le gisement éocène du Geiseltal (Allemagne, MP11 à MP14) a livré de nombreux fragments de chéloniens : plusieurs Trionychinae [12], le Testudininei (terrestre) Hadrianus eocenicus (Hummel, 1935) et des taxons de Testudinoidea d'eau douce, qui sont Borkenia germanica (Hummel, 1935) (groupe « Palaeochelys s. l. -Mauremys » [7,8]) et deux taxons du groupe des « Ptychogasteridae » : Geiselemys ptychogastroides (Hummel, 1935) et « Geoemyda » saxonica Hummel, 1935. Trois spécimens inédits du Geiseltal sont rapportés à un nouveau taxon de « Ptychogasteridae », permettant de définir ce groupe et d'identifier du matériel français. ...
... Le gisement éocène du Geiseltal (Allemagne, MP11 à MP14) a livré de nombreux fragments de chéloniens : plusieurs Trionychinae [12], le Testudininei (terrestre) Hadrianus eocenicus (Hummel, 1935) et des taxons de Testudinoidea d'eau douce, qui sont Borkenia germanica (Hummel, 1935) (groupe « Palaeochelys s. l. -Mauremys » [7,8]) et deux taxons du groupe des « Ptychogasteridae » : Geiselemys ptychogastroides (Hummel, 1935) et « Geoemyda » saxonica Hummel, 1935. Trois spécimens inédits du Geiseltal sont rapportés à un nouveau taxon de « Ptychogasteridae », permettant de définir ce groupe et d'identifier du matériel français. ...
... Le gisement éocène du Geiseltal (Allemagne, MP11 à MP14) a livré de nombreux fragments de chéloniens : plusieurs Trionychinae [12], le Testudininei (terrestre) Hadrianus eocenicus (Hummel, 1935) et des taxons de Testudinoidea d'eau douce, qui sont Borkenia germanica (Hummel, 1935) (groupe « Palaeochelys s. l. -Mauremys » [7,8]) et deux taxons du groupe des « Ptychogasteridae » : Geiselemys ptychogastroides (Hummel, 1935) et « Geoemyda » saxonica Hummel, 1935. Trois spécimens inédits du Geiseltal sont rapportés à un nouveau taxon de « Ptychogasteridae », permettant de définir ce groupe et d'identifier du matériel français. ...
Among the fossil turtles from of the Lutetian locality of Geiseltal (Germany), a new taxon of freshwater Testudinoidea is described. This taxon belongs to the ‘Ptychogasteridae’ group, which is diagnosed here. It is a primitive taxon, allowing the polarization of the characters of this group. A hypothesis of relationships between European and North American forms in the ‘Ptychogasteridae’ group is proposed. To cite this article: S. Hervet, C. R. Palevol 3 (2004).
... nov. sp., "Geoemyda" saxonica Hummel, 1935, from the Geiseltal Formation (Germany, Lutetian, MP11 and MP13), characters sampled from illustrations in Hummel (1935), Khosatzky and Młynarski (1966), and from GM-MLU 6986 (figured in Hummel, 1935, fig. 15) and GM-MLU 6985. ...
... nov. sp., "Geoemyda" saxonica Hummel, 1935, from the Geiseltal Formation (Germany, Lutetian, MP11 and MP13), characters sampled from illustrations in Hummel (1935), Khosatzky and Młynarski (1966), and from GM-MLU 6986 (figured in Hummel, 1935, fig. 15) and GM-MLU 6985. ...
... 15) and GM-MLU 6985. Geiselemys ptychogastroides (Hummel, 1935) from the Geiseltal Formation (Germany, Lutetian, MP11 and MP13); characters sampled from illustrations in Hummel (1935), Khosatzky and Młynarski (1966) and GM-MLU 6984. Hummelemys ambigua Hervet, 2004b from the Geiseltal Formation (Germany, Lutetian, MP12 and MP13), characters sampled from GM-MLU 10537, GM-MLU XXXV-221-1963, and GM-MLU XXXVI-45-1962, figured in Hervet, 2004b. ...
A new species of freshwater Testudinoidea is described from the fossil locality of Le Quesnoy (Oise, France; Argiles à Lignites du Soissonnais, Lower Eocene, Ypresian, MP7). This taxon is the oldest representative of Ptychogasteridae known from Europe. This species is compared to the American representatives of the genus Echmatemys Hay, 1906, to three Ptychogasteridae from the Geiseltal Formation (Eocene, Germany; Geiselemys ptychogastroides [Hummel, 1935], “Geoemyda” saxonica Hummel, 1935 and Hummelemys ambigua Hervet, 2004b), and to representatives of the Ptychogaster complex and to Clemmydopsis Boda, 1927. The phylogenetic relationships of this new taxon and of the other European Ptychogasteridae are studied.
... Although the species is, until now, exclusive to that locality, the genus has been recognised in the Ypresian to Bartonian records of several European countries. Thus, a second species is recognised, Pelorochelon eocaenica (Hummel 1935), defined by material from the Lutetian of Mücheln (Germany). The presence of Pelorochelon has not been confirmed in any other Iberian locality (Pérez-García 2017). ...
... Fish probably played a significant role in the diet of this species when available but based on its size and bite force, it is estimated that this crocodile would be able to prey on big vertebrates such as Lophiodon and Propalaeotherium during its adulthood (Hastings and Hellmund 2017). The tooth puncture mark present in the carapace now attributed to Pelorochelon eocaenica (Hummel 1935) specimen from the Geiseltal site (MP11) is considered compatible with the teeth morphology of Asiatosuchus, proving that these turtles could be a potential prey for these crocodiles as well (Hastings and Hellmund 2017). The specimens STUS-12064 and STUS-12065, from the MP13-14 levels of Casaseca de Campeán (Zamora) in the Duero Basin, correspond to a partial cranium and partial mandibles, respectively, and have been tentatively assigned to the genus Asiatosuchus (Martin de Jesús et al. 1987). ...
The Palaeogene sedimentary successions of the Duero Basin host an important record of fossil vertebrates, especially mammals and reptiles. The main fossil sites are placed at the west margin (Salamanca and Zamora areas) spanning from the Lutetian to the late Priabonian; and at the southeast tip (Almazán Basin) mainly Bartonian in age. The continued study of the sites is supplying new data that will improve the chronostratigraphy and the correlation between the western and eastern regions of the basin. The diversity and phylogenetic relationships of the represented fauna have been revised in recent years with the reassignment of some remains and the diagnosis of some new taxa. Four lineages of turtles are represented: Podocnemididae (Neochelys), Carettochelyidae (Allaeochelys), Trionychidae and Testudinidae (Pelorochelon). Crocodyliform paleobiodiversity is composed of notosuchians (close to Iberosuchus) and, at least, three crocodylian lineages: Planocraniidae (Duerosuchus), Alligatoroidea (Diplocynodon) and Crocodyloidea (‘Asiatosuchus’). The distribution of these taxa is not homogeneous throughout the basin, and a deep transformation is recognised throughout the middle Eocene. The fitting of the distribution and relationships of the reptilian fauna from the Duero Basin provides valuable information to understand some faunistic dynamics such as the endemism process of northwestern Iberia from the middle of the Eocene.
... Derived conditions: Several neural morphologies evolved from the primitive pattern: alternating octagonal and quadrangular neurals, or neurals with short postero-lateral sides. 1. Neurals with short postero-lateral sides appeared first in the Early to Middle Eocene in both Geoemydidae (Geoemyda saxonica and Geoemyda ptychogasteroides; Hummel, 1935) and Testudinidae (Hadrianus, Achilemys). 2. A clearly alternating pattern of octogonal and quadrangular neurals occurred in a clade of Testudinidae composed of Testudo, Indotestudo and Geochelone (s.l.). ...
... Phylogenetic hypotheses within this group vary considerably from one author to another, suggesting that homoplasy is rampant in this group (compare for example Hirayama, 1984, McCord et al., 2000, Yasukawa et al., 2001, Honda et al., 2002). The oldest species of this group may be "Geoemyda" ptychogasteroides from the middle Eocene of Germany ( Hummel, 1935) or "Cuvierochelys" parisiensis from the middle Eocene of France ( Botez, 1921;Hervet, 2004a). An unpublished new genus, formerly referred to Palaeochelys ( Nel et al., 1999) from the Early Eocene of France may be an older member of this family ( Hervet, 2003b). ...
... Von allen bis jetzt beschriebenen Hadrianus-Formen (vgl. DEBROIN 1977: 241) erinnert die besprochene Schildkröte vor allem an H. eocaenicus (HUMMEL, 1935). Sie kann auch mit Hadrianus spec, aus dem Eozän von Frankreich (DE BROIN 1977: 360, Fig. 102) verglichen werden. ...
... Wie erwähnt wurde, scheint unsere Hadrianus-Schildkröte mit der gut bekannten Art aus dem Eozän des Geiseltales (Sachsen, DDR)-Hadrianus eocaenicus-morphologisch "verwandt" zu sein (vgl. HUMMEL 1935, ZIMMERMANN-ROLLIUS 1967, AUFFENBERG 1974, MtYNARSKi 1977und DE BROIN 1977. Schon HUMMEL (op. ...
Mit 2 Abbildungen und 7 Tafeln) Manuskript eingelangt am 9. April 1985 Zusammenfassung In den pontischen Sand-und Schotterablagerungen von Prottes (Niederösterreich) konnte eine faunistisch und ökologisch bemerkenswerte Schildkröten-Fauna geborgen werden. Im systematischen Teil werden diese drei verschiedenen Schildkröten der Familie Testudinidae zugeordnet: 1. Testudo cf. promarginata REINACH, 1900, die durch einen verlängerten, glatten Panzer charakterisiert ist. 2. Testudo spec, mittelgroßes Tier mit stark skulpturiertem und gewölbtem Carapax ohne Cervicalschild (Merkmal der Gattung Geochelone). 3. Hadrianus spec, eine sehr große Schildkrötenart (Panzerlänge ca. 1 m) mit für diese Gattung sehr bezeichnenden, schaufelartigen Epiplastralplatten, die bis jetzt nur aus dem Eozän bekannt war. Diese drei Schildkröten repräsentieren drei verschiedene Entwicklungstendenzen und bewohnten drei verschiedene ökologische Nischen. Das in der Fundstelle Prottes in den vorzeitlichen Flußablage-rungen zusammengekommene Schildkrötenmaterial hatte einen refugialen und einen endemischen Charakter und weist auf ein lokal heißes, teilweise trockenes Klima hin. Abstract In the Pontian sand and gravel deposits of Prottes (Lower Austria) a turtle fauna has been collected which is remarkable in a faunistic and ecologie respect. In the systematical part of this paper these three different turtles are classified into the family Testudinidae: 1. Testudo cf. promarginata REINACH, 1900, which is characterized by a smooth and prolongated carapace. 2. Testudo spec., an animal of middle size with a strongly sculptured and vaulted carapace without a cervical shield (characteristic of the genus Geochelone). 3. Hadrianus spec, a very large turtle species (length of th carapace approx. 1 m) with shovel-like plates of the epiplastron. Hitherto this genus is only known from the Eocene. These three turtles represent three different trends of evolution and inhabited three different ecological niches. This material of turtles occuring in the river deposits of Prottes, has a refuge-like and endemic character of a local warm and partly dry climate.
... Autre espèce attribuée au genre : Ocadia ? germanica Hummel, 1935 =Borkenia germanica (Hummel, 1935) n. comb. ...
... Borkenia germanica (Hummel, 1935) n. comb. ...
Among fossil chelonians from the Tertiary of Western Europe, freshwater Testudinoidea (sensu Hirayama, 1985; Gaffney and Meylan, 1988) are abundant, and more particularly the “Palaeochelys sensu lato—Mauremys” group. This group has never been studied as a whole. Taxa of this group were before referred to extinct genera such as Ocadia, Clemmys, Emys or to the fossil genera Palaeochelys, Borkenia and Palaeoemys. These taxa are redefined, new taxa are described and a hypothesis of phylogenetic relationships is given.
... (Hadrianus) vialovi most closely resembles the European species M. (Hadrianus) eocaenica (Hummel, 1935) from the German middle Eocene (Hummel, 1935 ), M. (Hadrianus) cf. eocaenica from the Austrian middle Eocene (Bachmayer and Mlynarski, 1985b), and M. (Hadrianus) sp. ...
... (Hadrianus) vialovi most closely resembles the European species M. (Hadrianus) eocaenica (Hummel, 1935) from the German middle Eocene (Hummel, 1935 ), M. (Hadrianus) cf. eocaenica from the Austrian middle Eocene (Bachmayer and Mlynarski, 1985b), and M. (Hadrianus) sp. ...
Based on new material from the Andarak 2 locality in southern Kirghizia (early Eocene, late Ypresian), the testudinid Ergilemys vialovi Ckhikvadze, 1984, is assigned to the subgenus Hadrianus Cope, 1872, of the genus Manouria Gray, 1852. Part of the humerus of the oldest known cheloniine sea turtle is described from the same locality. The cheloniine sea turtles might have evolved during the Ypresian, an interval during which Cenozoic sea turtles were exceptionally diverse.
... The primitive members of the Ptychogaster group have acquired a neural irregularity, principally with neurals short sided posteriorly, rounded borders of the plastral lobes, dorsal thickening at gular-humeral boundary of the moderately elongated epiplastral lip, and an anterior position of the humeropectoral sulcus crossing the entoplastron. Among them, Geiselemys ptychogastroides (Hummel, 1935) (the holotype of Geoemyda saxonica not included), from the Lutetian (MP 13-14) of the Geiseltal, Germany, might be considered as the sister group of Ptychogaster, but it is not well described and poorly figured, although some reconsiderations were made (Hummel, 1935;Khozatsky and Mlynarski, 1966;Zimmermann-Rollius, 1966). Among other characters, the anterior border of the plastron is clearly concave, medially protruding in the juvenile. ...
... The primitive members of the Ptychogaster group have acquired a neural irregularity, principally with neurals short sided posteriorly, rounded borders of the plastral lobes, dorsal thickening at gular-humeral boundary of the moderately elongated epiplastral lip, and an anterior position of the humeropectoral sulcus crossing the entoplastron. Among them, Geiselemys ptychogastroides (Hummel, 1935) (the holotype of Geoemyda saxonica not included), from the Lutetian (MP 13-14) of the Geiseltal, Germany, might be considered as the sister group of Ptychogaster, but it is not well described and poorly figured, although some reconsiderations were made (Hummel, 1935;Khozatsky and Mlynarski, 1966;Zimmermann-Rollius, 1966). Among other characters, the anterior border of the plastron is clearly concave, medially protruding in the juvenile. ...
The Lower Miocene deposits of the Bardenas Reales of Navarre (NW Ebro Basin, northern Iberian Peninsula) have yielded a diverse vertebrate fauna, including remains of amphibians and reptiles. These remains occur in several localities in the Tudela Formation. The fossiliferous levels belong to the Biozones MN2b-3 (Biozones Z-A of the Ramblian, i.e., Late Aquitanian to Early Burdigalian in age). The amphibians and reptiles represent at least 13 out of 37 vertebrate species. Amphibians consist of a salamandrid urodele and two or three anurans. All the turtles are cryptodirans and consist of the chelydrid Chelydropsis apellanizi, the testudinids Ptychogaster (Temnoclemmys) bardenensis and Ptychogaster ronheimensis, and a Trionychinae indet. Squamates are represented by the anguid lizard Ophisaurus sp., a non-anguid lacertilian, an amphisbaenian, the erycine boid? Eryx sp., and indeterminate colubrids. Crocodilian remains are assigned to the basal alligatoroid Diplocynodon sp. The fossil associations of the Bardenas Reales of Navarre suggest that the vertebrates lived in the centre of an endoreic basin with stretches of water under intertropical to subtropical climatic conditions.
... The primitive members of the Ptychogaster group have acquired a neural irregularity, principally with neurals short sided posteriorly, rounded borders of the plastral lobes, dorsal thickening at gular–humeral boundary of the moderately elongated epiplastral lip, and an anterior position of the humeropectoral sulcus crossing the entoplastron. Among them, Geiselemys ptychogastroides (Hummel, 1935 ) (the holotype of Geoemyda saxonica not included), from the Lutetian (MP 13-14) of the Geiseltal, Germany, might be considered as the sister group of Ptychogaster, but it is not well described and poorly figured, although some reconsiderations were made (Hummel, 1935; Khozatsky and Mlynarski, 1966; Zimmermann-Rollius, 1966 ). Among other characters, the anterior border of the plastron is clearly concave, medially protruding in the juvenile. ...
... The primitive members of the Ptychogaster group have acquired a neural irregularity, principally with neurals short sided posteriorly, rounded borders of the plastral lobes, dorsal thickening at gular–humeral boundary of the moderately elongated epiplastral lip, and an anterior position of the humeropectoral sulcus crossing the entoplastron. Among them, Geiselemys ptychogastroides (Hummel, 1935 ) (the holotype of Geoemyda saxonica not included), from the Lutetian (MP 13-14) of the Geiseltal, Germany, might be considered as the sister group of Ptychogaster, but it is not well described and poorly figured, although some reconsiderations were made (Hummel, 1935; Khozatsky and Mlynarski, 1966; Zimmermann-Rollius, 1966 ). Among other characters, the anterior border of the plastron is clearly concave, medially protruding in the juvenile. ...
... X 19). Grâce à une variabilité reconnue, un type de décoration caractéristique peut se dessiner pour une espèce donnée, en l'occurrence pour (Reinach, 1900) in Cadena (2016)], ont (toujours en considérant les adultes) des pleurales à crêtes (plutôt que des bourrelets) plus fines que les bourrelets de T. michauxi, pour des cupules relativement plus grandes, de même que chez bien d'autres espèces : espèces soit indéterminées, soit définies par leur palais et mandibule, telles celle du Thanétien du Mont-Berru ou celles du Miocène (Broin, 1977 ;Owen & Bell 1849 ;Reinach, 1900 ;Hummel, 1935). L'absence de crâne à Rouzilhac ne permet pas une attribution définitive, mais la présence de cette décoration particulière de T. michauxi montre la similitude. ...
... Forachelys woodi was recovered nested within the Ptychogastrini and sister to the taxon saxonica (Fig. 14), which may be a junior synonym of Geiselemys ptychogastroides (Zimmermann-Rollius, 1966;Mlynarski, 1977;Hervet, 2006). This result is not unexpected considering similarities that F. woodi shares with G. ptychogastroides (Hummel, 1935) and the Ptychogaster/Temnoclemmys group. The epiplastra of F. woodi are most similar to numerous species of Ptychogaster, G. ptychogastroides and Bridgeremys pusilla. ...
Forachelys woodi gen. et sp. nov. is described from lower Miocene (Arikareean NALMA) exposures of the upper Las Cascadas Formation (~21 Ma) in the Panama Canal Basin. It represents the latest occurrence of ptychogastrine geoemydids in North America, a group known primarily from the early Eocene to late Miocene of Europe. Forachelys woodi shares features with taxa such as Geiselemys ptychogastroides (middle Eocene, Germany) and some species of the Ptychogaster/Temnoclemmys group (late Eocene to
late Miocene, Europe). The epiplastral morphology is furthermore comparable to Bridgeremys pusilla (middle Eocene of Wyoming and Utah), a taxon previously interpreted as a rhinoclemmydine ancestor. Within the Las Cascadas Formation, F. woodi co-occurred with an undescribed species of rhinoclemmydine geoemydid that represents the oldest record of Rhinoclemmys (sensu lato). The Miocene fossil record of testudinoids in Central America is reviewed and reexamined. Testudinids are diverse in the Miocene of Panama and Honduras, represented by aff. Stylemys, Caudochelys (2–3 large to giant species), and an undescribed genus and species that resembles ‘Testudo’ costarricensis. Rhinoclemmydines persisted in Panama almost continuously from the early to late Miocene, with records from the Las Cascadas (~21 Ma), upper Culebra (~20–19 Ma), Cucaracha (~19 Ma), and lower Gatun (~12–11 Ma) formations. At least two concurrent rhinoclemmydines are present in the Cucaracha Formation of Panama, and two
possibly concurrent taxa in the Gracias Formation (9–6.6 Ma) of Honduras.
... The taxonomy of large and giant fossil tortoise species has been puzzling for decades, and only recently have we started to understand their true diversity and their relationships. Historically, all giant tortoise species were considered as members of Testudo Linnaeus, 1758 (a name now restricted to five extant species and their closely related, extinct taxa; TTWG, 2021), like the majority of other tortoises, and were later transferred into other 'wastebasket genera ', in particular Cheirogaster Bergounioux, 1935and Geochelone Fitzinger, 1835(Bergounioux, 1935Hummel, 1935;Kuhn, 1964;Auffenberg, 1974;de Broin, 1977;Fritz & Bininda-Emonds, 2007;Georgalis & Kear, 2013;Luján et al., 2014;Vlachos et al., 2014). ...
Archaeological investigations carried out in the cave Zubbio di Cozzo San Pietro, Bagheria, Sicily, revealed the presence of a few skeletal elements of a large-sized tortoise in a funerary area dating to the Copper/Bronze Age. The tortoise has been AMS-dated revealing an age of 12.5 ± 0.5 kyr BP and therefore it pre-dates the funerary activities. The morphology of the retrieved skeletal elements differs from that of the only native tortoise currently living in Sicily, Testudo hermanni. The tortoise's size significantly exceeds the size range of extant Te. hermanni and all Testudo spp., as well as that of their known fossils, and suggests a shell length of 50-60 cm. Repeated efforts to obtain DNA sequences from the tortoise of Zubbio di Cozzo San Pietro failed, but the morphology of the femur is distinct enough to allow us to erect a new taxon, Solitudo sicula gen. et sp. nov., based on a parsimony analysis. It belongs to a hitherto unrecognized clade that includes other large-sized tortoises from Mediterranean islands, like Malta and Menorca. A review of the pertinent taxa indicates that the remains here described represent the geologically youngest large-sized tortoise of the Mediterranean area.
... Species of Echmatemys and Bridgeremys pusilla (Hay, 1908;Hutchison, 2006) have a simpler shaped ovoid four to five-sided N1. Ptychogastrines also have an ovoid N1, but this condition may be polymorphic in the middle Eocene taxon Geiselemys ptychogastroides (Hummel 1935). In UF 527000, N1-N8 are wider than long. ...
ABSTRACT Notapachemys oglala, gen. et sp. nov., is described based on carapace and plastron fossils collected from upper Eocene (Chadronian NALMA, White River Group) exposures in northwestern Nebraska, U.S.A. It is recognized as having a thick, smooth, partly fused, acarinate, egg-shaped carapace with strong plastral hindlobe kinesis. Many of these traits are present in some ptychogastrine geoemydids, a group known from the early Eocene to late Miocene of Europe. However, other aspects of shell morphology such as a six-sided Neural 1, medial gular notch, lack of distal gular tubercles, absence of inguinal scutes, and elongate kinetic inguinal buttresses resemble semi-aquatic to semi-terrestrial species of the New World tropically distributed genus Rhinoclemmys. The new taxon is one of the oldest plausible stem rhinoclemmydine geoemydids that possesses a mosaic of ptychogastrine and rhinoclemmydine features, suggesting the possibility of a close but unresolved relationship between these two groups. A convoluted history of ptychogastrines and rhinoclemmydines in North America is further exemplified in the early Miocene of the Panama Canal Basin, where ptychogastrine fossils from the Las Cascadas Formation (~22-20 Ma) and the oldest Rhinoclemmys fossils from the Cucaracha Formation (~18-16 Ma) have been collected in a region where Rhinoclemmys still lives today.
... The excavation campaigns and research led to a representative overview of the Geiseltal vertebrate taxa in the permanent exhibition (e.g. Krumbiegel, 1959aKrumbiegel, , 1959bKrumbiegel, , 1962Krumbiegel et al., 1983;Haubold, 1995). ...
The Geiseltal was a productive area for mining of lignite (brown coal) for about 100 years in central Germany (state of Saxony-Anhalt). Recognition of the scientific value of its famous fossil content came about in the 1920s, and from the early 1930s onwards Geiseltal is known as a unique Eocene terrestrial/palustrial Fossillagerstätte. During your professorship and position as Head of the geological Institute in Halle (Saale) in the 1920s, he focused more and more on the famous and outstanding finds of fossils in the middle Eocene brown coal of the Geiseltal. Ben Barnes was the pioneer of systematic and quantitative vertebrate excavations in the Geiseltal lignites. With his successful work, undertaken using with modern scientific aspects of his time, he gave rise for many other researchers to do so in the same way and to contribute step by step in reconstructing the geological and palaeontological history of fauna, flora and palaeoenvironment etc. in the Geiseltal. © 2018, Universidade Federal do Rio de Janeiro. All rights reserved.
... southern Europe is currently only part of the distribution area of small forms, but large taxa also lived there in warmer Cenozoic periods; see van Dijk et al., 2014). Thus, the European Paleogene record includes several forms of relatively large size, as the Eocene Pelorochelon eocaenica (Hummel, 1935) and Pelorochelon soriana Pérez-García et al., 2016, close to 70 cm, and the Oligocene Taraschelon gigas (Bravard, 1844), whose shell reached 80 cm. The favorable climatic conditions for this group of reptiles during the Cenozoic of Europe also enabled the development of taxa with larger sizes. ...
The presence of remains of a giant tortoise in the lower Pleistocene site of Fonelas P-1 (Guadix Basin, Betic Ranges; Granada, southeastern Spain) is reported and analyzed herein for the first time. This finding represents the youngest evidence of a large tortoise in continental Europe, dating the age of extinction of this successful lineage as several hundred thousand years younger than previously thought. So far, the most recent record known for continental Europe was at least 400,000 years older than the occurrence reported herein (Vaterá, Greece), that for the Spanish record being about 1.3 million years older (Las Higueruelas). This finding is justified as the youngest evidence of Titanochelon, a genus recorded in Europe since the beginning of the Miocene, which includes the largest terrestrial turtles known for the entire European fossil record. The decrease in the biogeographical distribution area and the final extinction of these temperature-sensitive animals in Europe is here recognized as a result of the climate changes documented during the Pliocene and lower Pleistocene. The identification of taxa with environmental and ecological requirements as restrictive as those known for the extant and extinct large tortoises, living in continental Europe 2.0 Ma, has important consequences. Thus, although the paleoclimatic inferences generally assumed for the whole of Europe interpret cooler and drier conditions at the end of the Pliocene, by the increase of the seasonality and the beginning of the glacial activity in the Northern Hemisphere, the record of Fonelas P-1 indicates that, in southern Europe or, at least, in the endorheic basins of the Betic Ranges, warmer climatic conditions than in the rest of the continent continued being present in these chronologies of the lower Pleistocene, being favorable for the persistence of these large tortoises.
... The material from Charneca do Lumiar corresponds to a shell of a single individual. The material is highly fragmented, taxa of Pan-Testudinidae recognized in the European record: the Eocene Fontainechelon cassouleti (Claude & Tong 2004), Pelorochelon eocaenica (Hummel 1935), Cheirogaster maurini and Pelorochelon soriana Pérez-García, Ortega and Jiménez Fuentes, 2016; the Oligocene Taraschelon gigas (Bravard 1844) and 'Ergilemys' bruneti Broin 1977; and the representatives of the Neogene Titanochelon. ...
The information about the European giant fossil tortoises has been greatly increased in recent years, based on the description of new material and the revision of previously published specimens. A genus with a wide paleobiogeographic and stratigraphic distribution, Titanochelon, has recently been described, containing all large testudinids from the European Neogene record. Its type species, Titanochelon bolivari, was described in the Spanish record. The presence of this species outside this country had not been justified. In this paper we present and describe fossil material of large testudinids from several sites in the Lisbon District (Portugal), from lower (Quinta da Barbacena; MN4), middle (Charneca do Lumiar and Quinta da Farinheira; MN5) and upper (Aveiras de Baixo; MN9) Miocene levels. This study allows us to confirm the presence of Titanochelon in the Portuguese record and justify, for the first time, the presence of Titanochelon bolivari outside Spain. Furthermore we are able to discuss the geographical and temporal distribution of the large tortoises in the middle Miocene of Europe, a relevant period for the diversity and evolution of this clade.
... Turtles were suggested as a probable prey item of Allognathosuchus (Carpenter & Lindsey, 1980) due to their very blunt teeth and stout jaw. Turtles were present at this time at Geiseltal (Hummel, 1935) but, due to the small size of Allognathosuchus, adult turtles were likely too large. However, juvenile turtles may have been an occasional prey item. ...
The Geiseltal fossil collection from southern Sachsen-Anhalt Germany contains remarkably well-preserved fossils of middle Eocene age. These include several crocodylian skulls, representing at least four different genera with a fifth genus represented by two mandibular rami. As sites with this many crocodylian genera are unknown in modern ecosystems, it has been hypothesized that these crocodylians may have differences in habit as compared to living crocodylians. In order to test similarities between the Geiseltal crocodylians and extant species, an analysis was conducted using geometric morphometrics to quantify shape in crocodylian skulls of all living species (
n
= 218) and all well-preserved crocodylian skulls of the Geiseltal fauna (
n
= 28). A relative warps analysis was used to quantify and compare skull shape, revealing
Allognathosuchus
and
Boverisuchus
to be very distinct from each other as well as from
Asiatosuchus
and
Diplocynodon
. Overlap in shape alone exists between some
Diplocynodon
and some
Asiatosuchus
, but there was significant difference in adult size. When compared with extant crocodylians, three Geiseltal genera occupied distinctly non-modern morphospace in the first two relative warps axes. Comparison of the diets of living crocodylians with similarly shaped skulls was used to reconstruct the prey preferences of the Geiseltal crocodylians, revealing differences in specialization. During the middle Eocene high global temperatures, partitioning of prey preference may have allowed this group to attain its higher than usual diversity, reducing the amount of direct competition.
... It differs from all other ptychogasterines (sensu Hervet, 2006) in the six apomorphic features mentioned above in the diagnosis. It further differs from all other ptychogasterines except "Geoemyda" saxonica Hummel, 1935 andClemmydopsis Boda, 1927 in the position of sulci between marginal 12 and vertebral 5 on pygal-although this feature cannot be evaluated in Geiselemys Khosatzky and Młynarski, 1966. It also differs from other ptychogasterines except Clemmydopsis in the longer peripherals (sometimes with marked anterior expansions), the entoplastron longer than broad and with more rounded margins, and the markedly narrower anterior border of the plastral lobe. ...
The taxonomy of the freshwater turtle subgenus Ptychogaster (Temnoclemmys) (Testudines,Geoemydidae, Ptychogasterinae) is revised based on new and previously-published mate-rial from the Vallès-Penedès Basin (NE Iberian Peninsula). Additional remains of subgenusPtychogaster, first reported from this basin, are also described. Although the oldest remainsof Temnoclemmys are left unassigned, most of the available remains are attributed to its typespecies, Ptychogaster (Temnoclemmys) batalleri, originally described from the Late Miocene(MN9 and MN10) of this basin. The new material expands the chronostratigraphic rangeof P. batalleri in the Vallès-Penedès Basin back to the MN7 + 8 and shows that “Testudo”celonica is its junior synonym. Emended diagnoses for Ptychogaster and Temnoclemmys arealso provided.
La taxonomie du sous-genre de tortue d’eau douce Ptychogaster (Temnoclemmys) (Testudines, Geoemydidae, Ptychogasterinae) est révisée sur la base du matériel nouveau et déjà publié provenant du bassin de Vallès-Penedès (Nord-Est de la péninsule Ibérique). Des restes additionnels du sous-genre Ptychogaster, signalé pour la première fois dans ce bassin, sont également décrits. Bien que les restes les plus anciens de Temnoclemmys ne soient pas assignés, la plupart des restes disponibles sont attribués à son espèce type, Ptychogaster (Temnoclemmys) batalleri, décrite à l’origine dans le Miocène supérieur (MN9 et MN10) de ce bassin. Le nouveau matériel élargit le cadre chronostratigraphique de P. batalleri dans le bassin de Vallès-Penedès à MN7 + 8 et montre que « Testudo » celonica est son synonyme plus récent. Des diagnoses émendées sont également fournies pour Ptychogaster et Temnoclemmys.
... Der Schildkrötenrest gehört zu einem Vertreter aus der Familie der Testudinidae, diese gehören zu den Landschildkröten. Die taxonomische Nähe zu Testudo eocaenica Hummel aus dem Geiseltal (Hummel 1935, Krumbiegel et al. 1983) bleibt anhand der mäßigen Erhaltung spekulativ. Der Nagernachweis belegt terrestrische Habitate. ...
In the fi lling of a sinkhole within the Lower Muschelkalk at the limestone quarry of the cement plant Karsdorf, beside fossil rich Upper Muschelkalk, also Palaeogene fossil material was found. The Muschelkalk part of the fi lling consists of approximately 50 m of Upper Muschelkalk and contains fossil material from the tetractinella bed (Trochitenkalk facies) up to the cycloides bed. The Palaeogene sediments of the collapse mass are not decalcifi ed and include a carpofl ora, rests of molluscs and vertebrates. The carpological samples show a wide stratigraphic range. Some species seem to be of Lower Eocene age or older (Tinospora wikinsonii Chandler) and correspond with appropriate fl oras from England like the London Clay; others are present from Eocene to Miocene. Most of the samples belong to Nyssaceans, Vitaceans, Menispermaceans and submerse swamp and aquatic plants (Potamogeton, Stratiotes, Sparganium, Ceratophyllum). Most of the small and fragile bone fragments belong to amphibians but also a lot of reptile material like turtles (Testudinoidea) and crocodiles were found. Special attention refers to mammal and bird remains. Exemplary the oldest rodent record in Germany (Paramyidae) needs to be mentioned. The faunistic fossils indicate close relations to the stratigraphically and spatially similar Geiseltal. In particular the gastropods Australorbis, Galba and Carychiopsis indicate close relations with the gastropod faunas of Geiseltal. Therefore most of the faunistic fossil content suggests a stratigraphic classifi cation in the Middle Eocene, but especially the carpofl ora indicates the possibility that the subrosion has already started in the Lower Eocene or earlier. With our data it is possible to more exactly determine the period of leaching in the Rötsalinar and thus the development of the sinkhole. © 2011 E. Schweizerbart'sche Verlagsbuchhandlung, Stuttgart, Germany.
We describe two exceptionally preserved fossil snakes from the Eocene Konservat-Lagerstätte of Geiseltal, located in the state of Saxony-Anhalt, Germany. The two snake specimens, GMH LIX-3-1992 and GMH XXXVIII-20-1964, can be confidently identified as booids based on general morphology and were thus compared to other geographically and/or temporally close fossil booids. We found that GMH LIX-3-1992 is morphologically very similar to Eoconstrictor spinifer, also from Geiseltal, and to Eoconstrictor fischeri, from the middle Eocene of Messel, but differs from both in a number of cranial and vertebral features. Based on these differences we erect the new species Eoconstrictor barnesi sp. nov.; GMH XXXVIII-20-1964 is very similar to GMH LIX-3-1992 and the two differ only in features that are likely ontogenetic. Phylogenetic analyses of snakes using maximum parsimony and Bayesian inference on datasets inclusive of both morphological and molecular data consistently support a close affinity of E. barnesi to E. fischeri and E. spinifer. Our preferred phylogenetic hypothesis places the three species of Eoconstrictor in a clade that is sister to Neotropical Boidae, a result consistent with previous studies. The genus Eoconstrictor could provide an important calibration point for molecular clock studies of booids and snakes in general.
We describe a new species of a giant tortoise of the genus Titanochelon from the locality of Sandelzhausen in south Germany (MN5, Burdigalian/Langhian boundary, Early/Middle Miocene). The material comprises at least two different individuals, one of which is a male individual preserving large parts of the carapace and plastron and several appendicular elements. The second individual is quite fragmented, preserving parts of the bridge and the posterior rim of the carapace. The new species, Titanochelon schleichi sp. nov., is the first species of a giant tortoise named from Germany and allows reconstructing an important diversity and expansion of titanochelones in the Western Palaearctic during the earlier parts of the Neogene.
The Geiseltal biota is an Eocene lacustrine Konservat‐Lagerstätte in central Germany. Despite its rich fauna and flora (over 50 000 fossil vertebrates, insects and other invertebrates, plants and trace fossils) the taphonomy of the biota, and of the anurans in particular, is poorly understood. We analysed the skeletal taphonomy of 168 anurans, scoring each specimen for orientation, completeness and articulation. Most are partial skeletons, truncated during preparation. This process introduces measurable artefacts into the taphonomic data, which requires further study. We identified recurring taphonomic states, including disarticulation or loss of abdominal bones, loss of one or more limbs, extensive disarticulation, and extensive loss of elements across the body. Relatively rare states include: loss of distal limb elements only, scattered bones, and isolated, articulated limbs. These data inform a new taphonomic model for the Geiseltal anurans. Comparative analysis of patterns in completeness and articulation in these specimens and anurans from other Cenozoic Lagerstätten reveals repeated patterns in preservation in lacustrine‐hosted settings, such as proximal to distal trends in completeness, preferential loss of small peripheral bones, disarticulation of abdominal bones, and reorientation of major body regions. Taphonomic controls in lacustrine systems are surface water temperature, lake depth, vegetation, exposure to bottom currents, transport mechanisms, scavenging, decay regime, bone size and location in the skeleton; all ultimately controlled by palaeoclimate, lake physiography and hydrology, and anuran anatomy. A universal approach to quantitative analysis of skeletal taphonomy metrics will enable comparative testing of the relationship between preservation and palaeoenvironment for diverse fossil vertebrates.
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.
Herein, we describe Alatochelon myrteum gen. et sp. nov., a large tortoise from the post‐Messinian (lower Pliocene) of the area of Puerto de la Cadena (Region of Murcia), Spain. The new taxon cannot be attributed to Titanochelon, which represented the only lineage of large tortoises previously recognized in the Neogene record of Europe. Alatochelon myrteum shows African affinities, especially with the extant African spurred tortoise Centrochelys sulcata. Although close phylogenetic relationships have previously been recognized among some tortoises of both continents, the dispersal of this lineage had always been proposed as having occurred in only one direction: from Europe to Africa. The dispersal of the lineage including the new Spanish form and Centrochelys sulcata from Africa to Europe is proposed here. This proposal is compatible with those previously recognized for some lineages of mammals also found in Puerto de la Cadena, identified as African lineages that probably reached Europe during the Messinian Salinity Crisis event. An African origin is also proposed for the lineage of Titanochelon. Therefore, the two lineages of large derived testudinids (i.e. Geochelona) recognized in the European record experienced diachronic dispersal events from Africa to Europe: that to which Alatochelon belongs probably during the Messinian and the other much earlier, at the beginning of the Miocene or before.
Turtles of the clade Pan-Trionychidae have a rich fossil record in the Old World, ranging from the Early Cretaceous (Hauterivian) to the Holocene. The clade most probably originated in Asia during the Early Cretaceous but spread from there to the Americas and Europe by the Late Cretaceous, to India and Australia by the Eocene, and to Afro-Arabia by the Neogene. The presence of a single pan-cyclanorbine in the Late Cretaceous (Maastrichtian) of Asia provides a minimum estimate for the age of the trionychid crown. As preserved, diversity was relatively high in Asia during the Late Cretaceous, but the subsequent, strong decline is likely a preservational bias, as extant faunas are relatively rich, especially throughout Asia. The range of trionychids contracted southward in Europe over the course of the Neogene, and the group is now locally extirpated. The group is now similarly absent from Arabia and Australia. A taxonomic review of the 180 named
Old World taxa finds 42 nomina valida, 38 nomina invalida, 88 nomina dubia, 11 nomina nuda, and 1 nomen suppressum.
A new extinct species of the genus Mauremys (Testudines: Geoemydidae) is described on the basis of three fossils from the late Pleistocene deposit of Tomori Amaga Cave on Miyakojima Island of the Miyako Island Group, Southern Ryukyus, Japan. Of these fossils, two (the anterior half of the plastron and the nuchal) were previously tentatively identified as Mauremys mutica, an extant species whose distribution is currently confined to the Yaeyama Island Group within the Ryukyus. The turtle represented by these two specimens and another, previously unreported material (left third peripheral) actually most resembles M. japonica from mainland Japan and the Northern Ryukyus, and M. yabei from the middle Pleistocene of mainland Japan. The fossil turtle however, differs from the other two in exhibiting a medial length of the entoplastron greater than the interhyoplastron in ventral view, and in having a longitudinal groove medial to the gulo–humeral sulcus on the epiplastron evident in dorsal view. The present finding strengthens endemicity of the recent terrestrial fauna of the Miyako Island Group.
A new extinct species of the genus Mauremys (Testudines: Geoemydidae) is described on the basis of three fossils from the late Pleistocene deposit of Tomori Amaga Cave on Miyakojima Island of the Miyako Island Group, Southern Ryukyus, Japan. Of these fossils, two (the anterior half of the plastron and the nuchal) were previously tentatively identified as Mauremys mutica, an extant species whose distribution is currentlyconfinedtotheYaeyamaIslandGroupwithintheRyukyus.The turtle represented by these two specimens and another, previously unreported material (left third peripheral) actually most resembles M. japonica from mainland Japan and the Northern Ryukyus, and M. yabei from the middle PleistoceneofmainlandJapan.Thefossilturtlehowever,differsfromthe other two in exhibiting a medial length of the entoplastron greater than the interhyoplastron in ventral view, and in having a longitudinal groove medial to the gulo–humeral sulcus on the epiplastron evident in dorsal view. The presentfindingstrengthensendemicityoftherecentterrestrialfaunaofthe Miyako Island Group.
An geoemydid turtle (Testudines: Geoemydidae) from the Upper Oligocene of Enspel (Westerwald, Rhineland-Palatinate): Another evidence of upper Oligocene turtle Palaeomauremys mlynarski/ (HERVET & LAPPARENT DE BROIN, 2000) from Rott is presented from lacustrine deposits of rhe crater lake of Enspel (Westerwald, Rhineland-Palarinate). The morphological characteristics, the raxonomic position and rheir palaeobiologic consequences are discussed. For the so-called "Palaeochelys-Mauremys- group or complex" the c1ear taxonomie term Palaeochelyinae n. subfarn. is proposed. A detailed overview shows the currenr comparable taxa.
New Late Oligocene (Chattian) remains of the terrapins Palaeoemys hessiaca Schleich, 1994 and Palaeomauremys tuberculata (Portis, 1882), the soft-shelled turtles Allaeochelys parayrei Noulet, 1867 and Trionyx cf. triunguis Forskål, 1775, and the snapping turtle Chelydrasia decheni (H. V. Meyer, 1852) from lacustrine sediments of the Oberleichtersbach doline (Lower Franconia, Bavaria, Germany) are reported. The morphological features of these five species, their taxonomic position and their palaeobiological implications are discussed. The new Palaeomauremys material suggests that P. mlynarskii (Hervet & Lapparent de Broin, 2000) is a junior synonym of P. tuberculata (Portis, 1882).
A ·review is given of the history of European chelonians, with the phases of development of the group. It starts with the beginning of the order Chelonii, at the end of the Triassic times, and leads to the present time. Geographic and stratigraphic distributions of the genera and families in the world and more especially in Europe, are indicated. Taxonomy contains the type species od each genus and the type locality of each species. This work is partly a compilation of unverified data; it includes several comments made in connection with some problematic taxa.
Well preserved eggs and eggshell from the Middle Eocene lignite beds of the Geiseltal (Eastern Germany) can be assigned to crocodilians and birds on the basis of their histostructure. Because a specific assignment of the crocodilian eggshell is not yet possible, its parataxonomic similarity to Krokolithes wilsoni Hirsch 1985 has supported establishing the new parataxonomic family Krokolithidae, including the new species, Krokolithes helleri. The avian eggshell of the Geiseltal is subdivided into three types, representing both palaeognathous and neognathous birds. The one specimen with a palaeognathous morphotype, has been assigned to the new parataxonomic family Medioolithidae with the type genus and species Medioolithus geiseltalensis.The eggshell, although fragmented, is excellently preserved, displaying color lines and impressions of decayed organic fibers in both the crocodilian and bird eggshell. Fragmentation may be due to transport and modern day weathering.
An extinct tortoise known from the uppermost Pleistocene of the Ryukyu Islands is described as the new species, Manouria oyamai (Testudines: Testudinidae) based on the skull, lower jaw and some postcranial elements. The specimens of M. oyamai were newly collected from the latest Pleistocene fissure deposits on Okinawa and Tokunoshima Islands, Ryukyu Islands, Japan. This is the first fossil record of Manouria sensu stricto. Phylogenetic analysis shows that Manouria sensu stricto, comprising M. emys, M. impressa and M. oyamai, is monophyletic. Manouria oyamai flourished in the Central and South Ryukyus until the Late Pleistocene and became extinct there, along with other endemic terrestrial vertebrates, in the latest Pleistocene.
New Late Oligocene (Chattian) remains of the terrapins Palaeoemys hessiaca Schleich, 1994 and Palaeomauremys tuberculata (Portis, 1882), the soft-shelled turtles Allaeochelys parayrei Noulet, 1867 and Trionyx cf. triunguis Forskål, 1775, and the snapping turtle Chelydrasia decheni (H. V. Meyer, 1852) from lacus-trine sediments of the Oberleichtersbach doline (Lower Franconia, Bavaria, Germany) are reported. The morphological features of these five species, their taxonomic position and their palaeobiological implications are discussed. The new Palaeomauremys material suggests that P. mlynarskii (Hervet & Lapparent de Broin, 2000) is a junior synonym of P. tuberculata (Portis, 1882).
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