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Leucocephalon yuwonoi McCord, Iverson, & Boeadi 1995:311 Holotype: MZB 10295; paratypes MZB 10296; UF 97333-35. The MZB types may have been lost in a fire (McCord, pers. comm.)., Published as part of Iverson, John B., 2022, A review of Chelonian type specimens (order Testudines), pp. 1-85 in Megataxa 7 (1) on page 25, DOI: 10.11646/megataxa.7.1.1, http://zenodo.org/record/5964441

Rhinoclemmys rubida (Cope 1869:148) Syntypes (9 or 10): ANSP 285 and 337–341 (Malnate 1971:354); and USNM 45612-14 (Cochran 1961:228; Reynolds et al. 2007:21), although Cope only noted 4 specimens, numbered 264– 267, in the original description, and 265–267 correspond to the USNM specimens. The fourth is either lost or part of the ANSP series (Reynolds et al. 2007:21). Additional syntypes may exist in the MNHN (Roger Bour, pers.comm.). Rhinoclemmys mexicana Gray 1870:659 Syntypes (3): BMNH 1947.3.4.34 (formerly 1870.6.20.1) and 1947.3.5.60-61 (1871.2.7.45- 46). Rhinoclemmys rubida perixantha Mosimann & Rabb 1953:1 Holotype: UMMZ 80336; photographed in the original description, Pl. I, and reproduced in Smith & Smith 1980:947); paratypes (5), UMMZ 80335, 80337, 104333, 104334 (formerly MCZ 53278 according to Kluge 1984:80), and 104335 (Kluge 1984:80)., Published as part of Iverson, John B., 2022, A review of Chelonian type specimens (order Testudines), pp. 1-85 in Megataxa 7 (1) on page 31, DOI: 10.11646/megataxa.7.1.1, http://zenodo.org/record/5964441, {"references":["Malnate, E. V. (1971) A catalog of primary types in the herpetological collections of the Academy of Natural Sciences, Philadelphia (ANSP). Proceedings of the Academy of Natural Sciences of Philadelphia, 123 (9), 345 - 375.","Cochran, D. M. (1961) Type specimens of reptiles and amphibians in the United States National Museum. United States National Museum Bulletin, 220, 1 - 291. https: // doi. org / 10.5479 / si. 03629236.220","Reynolds, R. P., Gotte, S. W. & Ernst, C. H. (2007) Catalog of type specimens of Recent Crocodilia and Testudines in the National Museum of Natural History, Smithsonian Institution. Smithsonian Contributions to Zoology, 626, 1 - 49. https: // doi. org / 10.5479 / si. 00810282.626","Gray, J. E. (1870) Supplement to the Catalogue of Shield Reptiles in the Collection of the British Museum. Part I. Testudinata (Tortoises). British Museum, London, 120 pp. https: // doi. org / 10.5962 / bhl. title. 11307","Smith, H. M. & Smith, R. B. (1980 [\" 1979 \"]) Synopsis of the herptofauna of Mexico. Volume VI. Guide to Mexican turtles. Bibliographic addendum III. John Johnson, North Bennington, Vermont, 1044 pp. https: // doi. org / 10.2307 / 1444548","Kluge, A. (1984) Type-specimens of reptiles in the University of Michigan Museum of Zoology. Miscellaneous Publications of the Museum of Zoology University of Michigan, 167, 1 - 85."]}

Morenia petersi (Anderson 1879:761) Syntypes (4, according to Bour 2009b:39, but only three identified): ZMB 8865 ( Fritz et al. 1994:164, photographed therein, p. 164, Fig. 3; illustrated in the original description, Pls. LIX and LXXV, and reproduced in Bour 2009b:38 and Das 2009:14); ZSI 155 and 156 (Das et al, 1998:126; Das 2009:5), but apparently lost (Kundu et al. 2018:37); and a fourth unlocated syntype. ZMB 8865 designated lectotype by Fritz et al. (1994:164)., Published as part of Iverson, John B., 2022, A review of Chelonian type specimens (order Testudines), pp. 1-85 in Megataxa 7 (1) on page 29, DOI: 10.11646/megataxa.7.1.1, http://zenodo.org/record/5964441, {"references":["Bour, R. (2009 b) Type specimens of Emys ocellata Dumeril & Bibron, 1835 with notes on the species of Morenia Gray, 1870. Emys, 16 (2), 33 - 42.","Fritz, U., Obst, F. & Gunther, R. (1994) Kritischer Typen- Katalog der Schildkrotensammlung (Reptilia: Testudines) des Zoologischen Museums Berlin. Mitteilungen aus dem Museum fur Naturkunde in Berlin. Zoologisches Museum und Institut fur Spezielle Zoologie Berlin, 70 (1), 157 - 175. https: // doi. org / 10.1002 / mmnz. 19940700110","Das, I. (2009) The discovery of Indian turtles, with notes on publications, type localities and type repositories. ENVIS Bulletin, 12, 1 - 14.","Das, I., Dattagupta, B. & Gayen, N. C. (1998) History and catalogue of reptile types in the collection of the Zoological Survey of India, Calcutta. Journal of South Asian Natural History, 3 (2), 121 - 172.","Kundu, S., Kumar, V., Murthy, B. H. C. K & Chandra, K. (2018) Chelonian types of National Zoological Collections. Zoological Survey of India, Kolkata, 70 pp. https: // doi. org / 10.26515 / rzsi / v 118 / i 1 / 2018 / 122300"]}

Malayemys khoratensis Ihlow, Vamberger, Flecks, Hartmann, Cota, Makchai, Meewattana, Dawson, Kheng, Rödder, & Fritz 2016:16 Holotype: THNHM 25816 (photographed in the original description, p. 17, Fig. 6); paratypes (3), MTD 49150; THNHM 25999; and ZFMK 97198. Types possibly sampled genetically in the original description. Malayemys isan Sumantha, Brophy, Kunya, Wiboonatthapol, & Pauwels 2016:2 Holotype: THNHM 25609 (photographed in the original description, Pls, 1, 5, 7); paratypes (6), CUMZ 0.2321; PSUZC RT 716-17; QSMI 1395- 96; and ZMKU R 000318. Three paratypes also photographed in the original description (Pls. 2, 5, 6, 13)., Published as part of Iverson, John B., 2022, A review of Chelonian type specimens (order Testudines), pp. 1-85 in Megataxa 7 (1) on page 25, DOI: 10.11646/megataxa.7.1.1, http://zenodo.org/record/5964441

Geoclemys hamiltonii (Gray 1830:9) Syntypes (2): BMNH 1947.3.4.41 (presumably the specimen illustrated by Gray 1831b:Pl. IX [as Emys guttata], according to Smith, 1931:111); and OUM 8477 (Nowak-Kemp & Fritz 2010:11). King & Burke (1989:35) erroneously listed BMNH 1947.3.4.41 as holotype. Emys guttata Gray 1831:Pl. 76 Holotype: BMNH, possibly 1947.3.4.41; illustrated in the original description (Pl. 76; see Smith 1931:11). Emys piquotii Lesson 1831:120 Holotype: Not located; possibly MNHN 1235. Emys hamiltonoides t Falconer & Cautley in Lydekker 1880:21 Holotype: Not located. Melanochelys pictus Murray 1884:107 Holotype: UKZM unnumbered; not located. Clemmys palaeindica t Lydekker 1885:178 Syntypes (2): NHM(P) 39838 and 39840 (Lydekker 1889:105); illustrated in the original description (Pl. XXI, Figs. 1, 1a, 1b, 3, 3a). Geoclemys sivalensis t Tewari & Badam 1969:555 Holotype: PUM A/665; illustrated in the original description (Fig. 2)., Published as part of Iverson, John B., 2022, A review of Chelonian type specimens (order Testudines), pp. 1-85 in Megataxa 7 (1) on page 24, DOI: 10.11646/megataxa.7.1.1, http://zenodo.org/record/5964441, {"references":["Gray, J. E. (1831 b) Synopsis Reptilium; or Short Descriptions of the Species of Reptiles. Part I. - Cataphracta. Tortoises, Crocodiles, and Enaliosaurians. Treuttel, Wurz, and Co., London, 85 pp. [Published May 1831]. https: // doi. org / 10.5962 / bhl. title. 4697","Smith, M. A. (1931) The Fauna of British India, including Ceylon and Burma. Reptilia and Amphibia. Vol. I. Loricata, Testudines. Taylor and Francis, London, 185 pp. https: // doi. org / 10.2307 / 1436417","Nowak-kemp, M. & Fritz, U. (2010) Chelonian type specimens at the Oxford University Museum. Zootaxa, 2604, 1 - 19. https: // doi. org / 10.11646 / zootaxa. 2604.1.1","King, F. W. & Burke, R. L. (1989) Crocodilian, Tuatara, and Turtle Species of the World. A Taxonomic and Geographic Reference. Association of Systematics Collections, Washington, D. C., 216 pp.","Lydekker, R. (1885) Indian Tertiary and post-Tertiary Vertebrata. Siwalik and Nerbada Chelonia. Memoirs of the Geological Survey of India, Palaeontologia Indica, (10) 3, 155 - 208.","Lydekker, R. (1889) Catalogue of the Fossil Reptilia and Amphibia in the British Museum (Natural History). Part III Chelonia. British Museum of Natural History, London, 239 pp."]}

Mauremys japonica (Temminck & Schlegel 1838:139) Syntypes (at least 6): RMNH 3331, 3332 (illustrated in Schlegel 1844:Pl. 41), 3333 (2 specimens), 3334, and 6142. Several syntypes illustrated in Temminck & Schlegel (1834:Pls. VIII–IX and 1838:Pls. VIII–IX). RMNH 3331 and 3333 photographed in Hoogmoed et al. 2010:12–13, Fig. 6–7). In addition, two specimens in the BMNH may also be syntypical (Hoogmoed et al. 2010:14). MNHN 1954 was erroneously reported as syntypical by King & Burke (1989:42; repeated by Iverson 1992:142)., Published as part of Iverson, John B., 2022, A review of Chelonian type specimens (order Testudines), pp. 1-85 in Megataxa 7 (1) on page 26, DOI: 10.11646/megataxa.7.1.1, http://zenodo.org/record/5964441, {"references":["Schlegel, H. (1844) Abbildungen neuer oder unvollstandig bekannter Amphibien, nach der Natur oder dem Leben entworfen, herausgegeben und mit einem erlauternden Texte begleitet. Arnz, Dusseldorf, 141 pp.","Temminck, C. J. & Schlegel, H. (1834) Reptilia. I. Les Cheloniens. In: Siebold, P. F. de. Fauna Japonica, sive Descriptio animalium, quae in itinere per Japoniam, jussu et auspiciis superiorum, qui summum in India Batava Imperium tenent, suscepto, annis 1823 - 1830 colleget, notis observationibus et adumbrationibus illustravit. Vol. III. Lugduni Batavorum. J. G. La Lau, Leiden. 80 pp., pls. I - IX.","Hoogmoed, M. S., Gasso Miracle, M. E. & van den Hoek Ostende, L. W. (2010) Type specimens of recent and fossil Testudines and Crocodylia in the collections of the Netherlands Centre for Biodiversity Naturalis, Leiden, the Netherlands. Zoologische Mededelingen Leiden, 84, 1 - 41.","King, F. W. & Burke, R. L. (1989) Crocodilian, Tuatara, and Turtle Species of the World. A Taxonomic and Geographic Reference. Association of Systematics Collections, Washington, D. C., 216 pp.","Iverson, J. B. (1992) A Revised Checklist with Distribution Maps of the Turtles of the World. Privately printed, Richmond, Indiana, 363 pp."]}

Sacalia bealei (Gray 1831:71) Syntypes (2): BMNH 1947.3.4.33 and 1947.3.4.42., Published as part of Iverson, John B., 2022, A review of Chelonian type specimens (order Testudines), pp. 1-85 in Megataxa 7 (1) on page 32, DOI: 10.11646/megataxa.7.1.1, http://zenodo.org/record/5964441

Melanochelys trijuga edeniana (Theobald 1876:12) Syntypes (7): ZSI 830, 1010, 1011, 1018, 1097, 1369 and 2589, according to Annandale (1913:71); but ZSI 930, 1010, 1011, 1018, 1097, 1369, and 1371, according to Das et al. (1998: 127) and Kundu et al. (2018:39). ZSI 930, 1011, 1097, and 1369 photographed in Kundu et al. (2018; 43, 45–46). Note that ZSI 830 is also a syntype of Emys trijuga var. burmana. Emys trijuga var. burmana Anderson 1879: 723 Syntypes (number uncertain): ZSI 830 (Das et al. 1998:126; photographed in Kundu et al. 2018:43– 44), others lost (Das et al. 1998:126; Kundu et al. 2018:39). Geoemyda trijuga wiroti Reimann in Nutaphand 1979:177 Holotype: Not designated or located, but in the ZFMK, according to H.-D. Philippen and N. Das (pers. comm. to John Iverson); illustrated in the original description (Figs. 66–67)., Published as part of Iverson, John B., 2022, A review of Chelonian type specimens (order Testudines), pp. 1-85 in Megataxa 7 (1) on page 29, DOI: 10.11646/megataxa.7.1.1, http://zenodo.org/record/5964441, {"references":["Annandale, N. (1913) The tortoises of Chota Nagpur. Records of the Indian Museum, 9 (5), 63 - 78.","Das, I., Dattagupta, B. & Gayen, N. C. (1998) History and catalogue of reptile types in the collection of the Zoological Survey of India, Calcutta. Journal of South Asian Natural History, 3 (2), 121 - 172.","Kundu, S., Kumar, V., Murthy, B. H. C. K & Chandra, K. (2018) Chelonian types of National Zoological Collections. Zoological Survey of India, Kolkata, 70 pp. https: // doi. org / 10.26515 / rzsi / v 118 / i 1 / 2018 / 122300"]}

Herpesviruses are important pathogens in tortoises and turtles, yet little is known about the epidemiology of these viruses. We analyzed herpesviruses detected by PCR in samples from captive chelonians in Europe according to virus strain, host species, year and season in which the animal was tested, and country in which the animal was kept. A total of 4,797 samples submitted to a diagnostic laboratory in Europe between January 2016 and December 2020 were evaluated. Of these, 312 (6.50%) were positive for herpesviruses. The types most commonly found were testudinid herpesvirus (TeHV)1 (143 positive, 45.83%) and TeHV3 (153 positive, 49.04%), but also included TeHV2 (1 positive, 0.32%), TeHV4 (3 positive, 0.96%), Terrapene herpesvirus 1 (7 positive, 2.24%), Trachemys herpesvirus 1 (2 positive, 0.64%), and three previously undescribed herpesviruses (0.96%). Herpesviruses were detected in chelonians in the families Testudinidae, Emydidae, Geoemydidae, and in the suborder Pleurodira. Among the species for which 100 samples or more were available, the highest proportions of positive samples (positivity rates) were found in samples from Horsfield's tortoises (Testudo horsfieldii) (14.96%), and radiated tortoises (Astrochelys radiata) (14.05%). Among tortoises (Testudinidae), viruses were most often detected in the spring, while in emydid turtles (Emydidae) they were most often detected in the summer. A comparison of the positivity rates according to country showed significant differences, with the highest rate in samples from Italy (16.01%). This study indicated possible differences in herpesvirus positivity rates depending on host species, virus strain, year of sampling, season, and country of origin. It provides useful information in further understanding fluctuations in infection rates as well as in helping to guide decision making for herpesvirus diagnostics in chelonian patients. It also provides evidence for the international dispersal of herpesviruses with their hosts through international trade.

Background Quantity discrimination, the ability to discriminate a magnitude of difference or discrete numerical information, plays a key role in animal behavior. While quantitative ability has been well documented in fishes, birds, mammals, and even in previously unstudied invertebrates and amphibians, it is still poorly understood in reptiles and has never been tested in an aquatic turtle despite the fact that evidence is accumulating that reptiles possess cognitive skills and learning ability. To help address this deficiency in reptiles, we investigated the quantitative ability of an Asian freshwater turtle, Mauremys sinensis, using red cubes on a white background in a trained quantity discrimination task. While spontaneous quantity discrimination methods are thought to be more ecologically relevant, training animals on a quantity discrimination task allows more comparability across taxa. Results We assessed the turtles’ quantitative performance in a series of tests with increasing quantity ratios and numerosities. Surprisingly, the turtles were able to discriminate quantities of up to 9 versus 10 (ratio = 0.9), which shows a good quantitative ability that is comparable to some endotherms. Our results showed that the turtles’ quantitative performance followed Weber’s law, in which success rate decreased with increasing quantity ratio across a wide range of numerosities. Furthermore, the gradual improvement of their success rate across different experiments and phases suggested that the turtles possess learning ability. Conclusions Reptile quantitative ability has long been ignored and therefore is likely under-estimated. More comparative research on numerical cognition across a diversity of species will greatly contribute to a clearer understanding of quantitative ability in animals and whether it has evolved convergently in diverse taxa.

We present new fossil records of the geoemydid turtle Bridgeremys pusilla from the Uinta Formation of Utah. Turtles are abundant throughout the unit, and known taxa are similar to those from the older strata in the Upper Green River Basin in Wyoming from the Bridger and Washakie Formations. B. pusilla is known from Bridgerian deposits but was not previously known from after the Turtle Bluff Member of the Bridger Formation. The taxon was coveal with two species of the geoemydid Echmatemys (E. callopyge and E. wyomingensis), a common genus of extinct pond turtles known primarily from lacustrine and fluvial deposits in western North America, including the Uinta Basin. In addition to previously documented morphological differences, our geometric morphometric analyses revealed significant differences in epiplastral morphology between B. pusilla and the two coeval Echmatemys species. Bridgeremys pusilla shared several morphological characters with Testudinidae. However, our anatomical network analysis suggests that the carapace of B. pusilla distributed stress forces in a manner more similar to emydids (basal and derived) than to derived testudinoids (Testudinidae and Emydidae), including Echmatemys species. This finding changes our understanding of the ecology of the species and sheds light onto how geoemydid turtles of the Uinta Formation may have partitioned the available ecospace. These new Uintan records extend the geographic range of B. pusilla into the Uinta Basin and stratigraphically through the top of the Uinta Formation, extending the temporal range of the taxon by more than 4 million years through the Uintan North American Land Mammal Age to the base of the Duchesne River Formation.

An individual of the Assam Leaf Turtle Cyclemys gemeli was sighted at an elevation of 580m in Darjeeling. This is the first record of the species from the Darjeeling-Sikkim Himalaya, India. Photographs and locality record of the sighting are provided in the present communication.

Mauremys reevesii (Gray, 1831), which belongs to Mauremys of Geoemydidae (Testudines), distributed in China, as well as Japan and Korea. Previous studies have developed several polymorphic microsatellite loci, but most of them were dinucleotide motifs. Here, we developed 15 polynucleotide-repeat microsatellite loci (including di-, tri, tetra-and pentanucleotide motifs) for M. reevesii through Restriction-site Associated DNA tags sequencing (RAD-seq). A total of 987 microsatellite loci with flanking sequences were suitable for setting primers for polymerase chain reactions (PCR). To verify the identified SSRs, 40 primer pairs were selected for PCR detection. In total, 32 primer sets produced strong PCR products matching their expected sizes, in which species amplification tests showed that 15 were polymorphic. And the number of alleles per locus ranged from 3 to 16. The observed and expected heterozygosity per locus varied from 0.3784 to 1.000 and from 0.3995 to 0.9700, respectively. The methodology of microsatellite isolation constructed in this study is not only cost-effective and time-saving in comparison to traditional approaches, but also can be served as useful tools which benefit population genetics studies and conservation management of M. reevesii.

Falcaustra tintlwini sp. nov. (Ascaridida, Kathlaniidae) from the large intestine of Batagur trivittata (Testudines, Geoemydidae) is described and illustrated. Falcaustra tintlwini represents the 20th Oriental species assigned to the genus and is distinguished from other Oriental species by the distribution pattern of the caudal papillae (6 precloacal, 12 postcloacal, and 1 median), length of spicules (0.43-0.50 mm) and absence of a pseudosucker.

We report the southernmost distribution of Eastern Black-bridged Leaf Turtle Cyclemys pulchristriata Fritz, Gaulke & Lehr, 1997, in an isolated mountain in Ba Ria–Vung Tau Province, southern Vietnam based on morphological and molecular data of a wild juvenile captured in the wild. This record extends the distribution of this species about 150 km south of previous records and 600 km from its type locality.

This study collected records of 135 taxa of parasitic helminths (Nematoda, Trematoda, Cestoda, Monogenoidea and Acanthocephala) and ectosimbionts (Temnocephalida) associated with continental Testudines from South America. Eighty-nine helminths were identified at the species level while others were identified up to genus or family levels. The greatest diversity of helminths associated with Testudines was reported in Brazil. Chelidae was the family with the largest number of helminth species. Regarding the conservation status, 17 Testudines species with helminth records are cited in the IUCN Red List of Threatened Species. Knowledge of helminth biodiversity and of relations between these organisms and Testudines can provide important data on host biology. Therefore, information generated by studies of helminths can contribute to research which aims at the conservation of organims and their habitats. En este estudio se recogieron registros de 135 taxones de helmintos parásitos (Nematoda, Trematoda, Cestoda, Monogenoidea y Acanthocephala) y ectosimbiontes (Temnocephalida) asociados a Testudines continentales de Sudamérica. Se identificaron 89 helmintos a nivel de especie, mientras que otros se identificaron hasta el nivel de género o familia. La mayor diversidad de helmintos asociados a los Testudines se registró en Brasil. Chelidae fue la familia con el mayor número de especies de helmintos. En cuanto al estado de conservación, 17 especies de Testudines con registros de helmintos están citadas en la Lista Roja de Especies Amenazadas de la IUCN. El conocimiento de la biodiversidad de los helmintos y de las relaciones entre estos organismos y los Testudines puede aportar datos importantes sobre la biología de los hospedadores. Por lo tanto, la información generada por los estudios de los helmintos puede contribuir a la investigación que tiene como objetivo la conservación de los organismos y sus hábitats.

Testudinoidea represents an evolutionarily unique taxon comprising both turtles and tortoises. The contrasting habitats that turtles and tortoises inhabit are associated with unique physio-ecological challenges hence enable distinct adaptive evolutionary strategies. To comparatively understand the pattern and strength of Darwinian selection and physicochemical evolution in turtle and tortoise mitogenomes, we employed adaptive divergence and selection analyses. We evaluated changes in structural and biochemical properties, and codon models on the mitochondrial protein-coding genes (PCGs) among three turtles and a tortoise lineage. We used mitochondrial PCGs that constitute the crucial oxidative phosphorylation (OXPHOS) respiratory system, a critical metabolic regulator which assumes key significance in energy regulation of ectotherms.We detected strong evidence of positive selection along the turtle lineages: Geoemydidae, Emydidae, and Platysternidae, but relatively weak signals in tortoises. The Platysternidae turtles revealed the highest gene and site-wise positive selection. In turtles, positively selected sites were prevalent in NAD2 and NAD4 genes in OXPHOS Complex I, and COB gene of Complex III, indicating convergent adaptive evolution. Besides, NAD3 was the only subunit that showed adaptive selection in both turtles and tortoises, expressing its relevance for all Testudinoidea. Structural and functional analysis revealed many sites and physiochemical changes in important conserved as well as biomedically significant regions, suggesting the influence of adaptive pressure on mitogenome functions. Hence, our study furnished novel evidence of contrasting evolutionary selective pressure acting on closely related groups such as turtles and tortoises with unique habitat preferences and associated eco-physiological challenges.

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.

New Pleistocene fossilized turtle remains from five localities of western Ecuador (Santa Elena Province) are described here. All these shell (carapace and plastron) fossil remains come from the Tablazo Formation and belong to three different lineages of cryptodires (“hidden-necked” turtles). The most abundant remains belong to geoemydids, attributed here to the genus Rhinoclemmys (indeterminate species). Less abundant in occurrence are the kinosternidids, attributed to Kinosternon (indeterminate species), and the first fossil record of chelydrids, Chelydra(indeterminate species), in the entirety of Central and South America.

Altres ajuts: Forschungskredit of the University of Zurich (FK-20-110), Former postdoctoral grant from the University of Torino, Fondi di Ateneo dell'Università di Torino (2019-2020) We here describe a small turtle assemblage originating from the early Tortonian (late Miocene) palaeoisland of Scontrone, central Italy, a locality previously known mostly for its endemic mammals and giant birds, which were otherwise shared only with the Gargano localities, another fossiliferous area belonging to the same palaeobioprovince. The fossil turtle remains from Scontrone are referred to the geoemydid Mauremys sp. and a so far unidentified large-sized testudinid. The biogeographic origins of the Scontrone insular chelonians are discussed. The Scontrone geoemydid adds to the known occurrences of Mauremys in the late Miocene of the Mediterranean. The Scontrone large tortoise represents the oldest known Mediterranean insular testudinid, predating significantly the well-known Quaternary endemic island tortoises of the area.

Revealing the genetic basis of the existence of different species living together in different geographic regions provides clarification of this phylogeographic differentiation. In this study, we investigated the population genetics and evaluated the level of genetic variation of inland and coastal populations of Mauremys and Emys in Turkey. Tissue samples of 196 terrapins were studied which were collected from syntopic coastal (Golbent-Soke/Aydin; M. rivulata and E. orbicularis) and inland populations (Bahcesaray/Aksaray; M. caspica and E. orbicularis). DNA was isolated using the InnuPREP DNA Mini Kit. Mitochondrial DNA sequences and allelic variation at 13 microsatellite loci for Mauremys and 12 microsatellite loci for Emys were examined. Three haplotypes were found for Emys orbicularis (Im, Ip and Iw) collected from the coastal region and two haplotypes for Emys orbicularis (Ig and Im) collected from inland. Two haplotypes were identified for M. caspica (Cmt8 and Cmt9) and three haplotypes were identified for M. rivulata (Rmt3, Rmt24 and Rmt26). Using microsatellites and the software STRUCTURE the most probable value for K was revealed as two 2 for both species. The FST value between M. rivulata and M. caspica was 0.39, and between the coastal and inland populations of E. orbicularis 0.09. It can be concluded that Emys populations tend to evolve by somehow preserving the allelic richness they have and Mauremys populations continue to differentiate so that new species emerge in the evolutionary process to reach the ideal allelic structure.

A Late Pleistocene fossil represented by an incomplete right hypoplastron of Cuora amboinensis was recovered from the Wajak site (ca. 37 ka–29 ka: Late Pleistocene) in East Java, Indonesia. The fossil constitutes the first certain prehistoric record of this species from Java, which implies that the current conspecific population in this island is indigenous to the region rather than artificially introduced from the Southeast Asian continent in historic times. The specimen has four small distinct impact pits on the hypoplastron in dorsal view, which were possibly caused by a pointed stone artifact or a bone tool. The presence of such percussion marks suggests that this turtle was consumed by the Wajak people.

The marginal populations of the Sahara Blue-eyed pond turtle, Mauremys leprosa saharica (Testudines: Geoemydidae) in the pre-Saharan area in the Lower Draa valley, are facing extreme environmental conditions of arid climate and anthropogenic activities, including overuse of water and land salinization. In this study, we investigated a small isolated population of the Blue-eyed pond turtle at Sidi El Mehdaoui oasis, Lower Draa valley, to determine its population structure and morphometric characteristics. Water salinity was about 24% of seawater and dissolved oxygen concentration was less than 3 mg · L−1 (sub-hypoxia). Turtles were captured, sexed, weighed, and measured for the carapace dimensions before being released. The average body size of adults (carapace length) was much smaller than those in other Moroccan populations. Among adult individuals, the mean carapace lengths were 103 ± 21 mm and 104.4 ± 35 mm for males and females, respectively. The corresponding mean body masses were 173 ± 92.4 g and 135 ± 52 g. There were no significant differences between sexes for size or weight (P > 0.05). The studied population is small with a predominance of adults (75%). Sexual dimorphism was found to be significant in shape but not in size. In fact, the Sexual Dimorphism Index for size had the lowest level ever described for the species, possibly because of the low productivity of this marginal isolated habitat. The increased drought and salinization are great threats to the long-term persistence of the vulnerable populations of the Saharan blue-eyed pond turtle and their habitats. Therefore, conservation measures of these populations through the protection of their habitats are very urgent.

The Asian box turtle genus Cuora currently comprises 13 species with a wide distribution in Southeast Asia, including China and the islands of Indonesia and Philippines. The populations of these species are rapidly declining due to human pressure, including pollution, habitat loss, and harvesting for food consumption. Notably, the IUCN Red List identifies almost all species of the genus Cuora as Endangered (EN) or Critically Endangered (CR). In this study, we explore the karyotypes of 10 Cuora species with conventional (Giemsa staining, C-banding, karyogram reconstruction) and molecular cytogenetic methods (in situ hybridization with probes for rDNA loci and telomeric repeats). Our study reveals a diploid chromosome number of 2n = 52 chromosomes in all studied species, with karyotypes of similar chromosomal morphology. In all examined species, rDNA loci are detected at a single medium-sized chromosome pair and the telomeric repeats are restricted to the expected terminal position across all chromosomes. In contrast to a previous report, sex chromosomes are neither detected in Cuora galbinifrons nor in any other species. Therefore, we assume that these turtles have either environmental sex determination or genotypic sex determination with poorly differentiated sex chromosomes. The conservation of genome organization could explain the numerous observed cases of interspecific hybridization both within the genus Cuora and across geoemydid turtles.

The Siam Shield Leech, Placobdelloides siamensis, is a common leech found on Malayemys turtles in Thailand. Sixty Snail-eating Turtles (29 Malayemys macrocephala and 31 M. subtrijuga) were caught over twelve months (February 2017 – January 2018) to determine host characteristics (body size, weight and sex), parasitism (prevalence, intensity and density) and seasonal aquatic environmental factors (conductivity, nitrate nitrogen, dissolved oxygen, pH, salinity and total dissolved solids). There was no significant difference of infection rate between species and sex in both turtle species. Leech prevalence indicated that all turtle individuals were infected throughout year, while the infection rate was significantly higher in larger and heavier turtles mainly on the carapace with an average number of leech approximately 474.80 ± 331.38 individuals for individual host infection and 76.53 ± 20.27 individuals for infection per 100 g body weight. The high level of leech parasitism also caused a rot wound and shell hole which caused the host to die. Aquatic environmental factors did not influence the infection of leeches in both turtle species. Therefore, the factors that influenced the infection rate of P. siamensis were based on only host body size and weight without effect from season. In addition, this study also showed two new hosts, including Cyclemys oldhamii and Heosemys grandis and the widespread distribution from northern, north-eastern, western, central and southern Thailand were reported.

Batagur trivittata Dum��ril & Bibron, 1835 Our sample (N = 392) included neonate B. trivittata sourced from the captive-breeding group at YZG (N = 261) and hatched from eggs deposited by wild females along the Chindwin River and head-started at LV (N = 131; Fig. 1). Neo- nates at both sites were examined and measured 12���36 hours after hatching. Using a pair of dial calipers we measured (to the nearest 1.0 mm) CL (Method D of Iverson & Lewis 2018), maximum carapace width (CW), mid-line plastron length (PL; from base of anal notch to anterior margin of gular scute), and shell depth (SD; from plastron to highest point of carapace) of each neonate. We used a Pesola�� spring scale (�� 5.0 g) and Camry�� digital balance (�� 0.001 g) to determine body mass (BM) of neonates at YZG and LV, respectively. We photographed neonates with a Canon 7D EOS Mark II (with Canon Macro lens EF 100 mm 1:2.8 USM) and Sony A6000 digital cameras. We made comparisons with older cohorts being head-started at LV to document the ontogeny of certain shell attributes. We obtained measurements of shell attributes from all neonates in our sample, but report values for body mass only for neonates at LV because the wide margin of error associated with spring scales used at YZG precluded a sufficiently precise measure of this attribute. Mean �� 1SD and range (in parentheses) of shell attributes and body mass of neonate B. trivittata are as follows: CL = 55.3 �� 4.0 mm (43.0���65.0 mm); CW = 51.7 �� 4.6 mm (35.0��� 61.8 mm); PL = 49.4 �� 4.3 mm (35.0���59.0 mm); SD = 30.4 �� 3.7 mm (20.0���48.0 mm); BM = 47.7 �� 4.4 mm (36.3���60.0 mm). Assuming an upper asymptotic CL of 580���620 mm for adult female B. trivittata (Smith 1931; Platt et al. 2019), the mean CL of neonates is 8.9���9.5% of the maximum adult body size. Mean neonate body mass was 74.7% of mean egg mass for eggs collected during the 2017���18 and 2018���19 nesting seasons (mean egg mass = 63.8 �� 4.0 g; range = 50.3���70.0 g; N = 137; Platt et al. unpubl. data). In neonate B. trivittata the overall shape of the carapace is ovoid to almost circular (mean CW = 93% mean CL), being widest at the third vertebral and sixth marginal pair, and tapering posteriorly (Fig. 2). The carapace is green to brown with dark elongated blotches that straddle adjacent pleural scutes 1���4. The terminal blotch on the fourth pleural scute does not extend onto the adjacent fifth vertebral, but instead is present at the junction of the pleural, vertebral, and marginal scutes, and much reduced. Similar dark pleural blotches are also present in hatchling B. borneoensis and B. dhongoka, but absent in B. affinis, B. baska and B. kachuga (Moll 1986; Praschag et al. 2009; Moll et al. 2015; Guntoro 2017; Turtle Survival Alliance, unpublished photographs). Notably, molecular analyses indicate that B. borneoensis and B. dhongoka are successive sister taxa to B. trivittata, while B. affinis, B. baska, and B. kachuga constitute a separate clade of Batagur (Minh et al. 2007; Praschag et al. 2007, 2009). Thus, patterns of carapace coloration among hatchings appear to reflect the underlying phylogenetic relationships in Batagur. The medial keel is well-pronounced on vertebrals 1���4 with sharp, spine-like posterior-pointing projections on vertebrals 2���4. The posterior edge of marginals 1���7 are tipped with a sharp rearward-pointing spine that becomes increasingly larger, reaching its greatest development on marginal 7. Marginal 6 is weakly serrated; marginals 7���12 are deeply serrated with the posterior edge of each scute (except 12) curving rearwards. Faint grey-black elongated blotches are present on marginals 4���10, overlapping the intermarginal seams; these fade rapidly and are no longer evident within a few weeks of hatching. The plastron, bridge, and ventral surface of the marginals are whitish to cream. An umbilical scar is present on the mid-line of the abdominal scute of very young neonates, but disappears rapidly with age (Fig. 3A). A prominent keel-like bony ridge extends along the lateral edge of the plastron from the humeral to anal scutes, forming a distinct angle between the plastron and bridge regions of the pectoral and abdominal scutes. The posterior edge of this ridge on each scute is tipped with a sharp rearward projecting spine. This spine is well-developed on the pectoral, abdominal, and femoral scutes; less so on the anal scute, and weakly developed on the humeral scute (Fig. 3B). Soft parts of neonates vary in coloration from uniform green-brown to dark grey dorsally, with most individuals tending towards the latter; the ventral surface of the legs, tail and gular region are lighter (white-yellowish). The rostrum extends horizontally from the snout with no upturn. The keratinized sheath of the maxilla is slightly serrated while that of the mandible is smooth; a caruncle (egg-tooth) is present just below the rostrum in very young neonates (Fig. 4). Neonates hatched in mid- to late May at LV retained the caruncle through June, but the structure was no longer present in any individual at the end of July. The fore- and hind-feet are fully webbed with sharp claws (five and four on each fore- and hind-foot, respectively) extending just beyond the light-colored margin. The iris is greenish-yellow with a dark pupil. A number of notable ontogenetic changes occur as neonate B. trivittata mature. The dark elongated blotches on the pleural scutes of neonates are absent from older females, which have a solid dark brown-gray carapace (Smith 1931; Platt et al. 2017b). In adult males, the pleural blotches probably coalesce to form the black lateral stripes on the pleural scutes (Smith 1931), although we have not yet monitored cohorts long enough to confirm the origin of these stripes. The medial carapace keel is present but lower and less pronounced in adults. Our observations of successive cohorts being headstarted at LV indicate the vertebral spines are blunt after two years of growth, with the exception of the spine on vertebral 4, which remains pointed and sharp to the touch until 3���4 years of age. The rearward pointing spines on marginals 1���7 remain sharp to the touch in one-year-old turtles, become less so in two-year-olds, and are blunt and rounded by age three. Marginals 7���12 become progressively less serrated as turtles mature, with the posterior margin of the carapace becoming smooth by age five. Coloration of the plastron, plastral bridge, and ventral surface of the marginals in females change from whitish-cream to dark brown-black with increasing age and by age five these surfaces are heavily pigmented (Platt et al. 2017b). In contrast, the plastron, plastral bridge, and ventral surface of the marginals in males remain whitish-cream into maturity (Platt et al. 2017b). The bony ridge extending along the distal edge of the plastron becomes smoother with age and is no longer evident in five-year-old B. trivittata. The spines adorning this ridge remain sharp to the touch in two-year-old turtles, become blunt by age three, and have largely disappeared in four-year-olds. The functional significance of the spines on the vertebrals and plastral ridge of neonate B. trivittata is unknown; however, the spines on other turtles are generally interpreted as an important physical defense that deters consumption by gape-limited predators such as large fish and wading birds (Inbar & Lev-Yadun 2005). The fact that spines on B. trivittata disappear as the turtle matures and body size increasingly provides safety from predation seems consistent with the anti-predator defense hypothesis., Published as part of Platt, Steven G., Lwin, Tint, Win, Myo Min, Platt, Kalyar, Haislip, Nathan A., Dijk, Peter Paul Van & Rainwater, Thomas R., 2020, First description of neonate Batagur trivittata (Testudines: Geoemydidae), pp. 394-400 in Zootaxa 4821 (2) on pages 394-399, DOI: 10.11646/zootaxa.4821.2.10, http://zenodo.org/record/4015089, {"references":["Dumeril, A. M. C. & Bibron, G. (1835) Erpetologie generale ou histoire naturelle complete du reptiles. Volume II. Chelonians. Roret, Paris, 680 pp. https: // doi. org / 10.5962 / bhl. title. 45973","Iverson, J. B. & Lewis, E. L. (2018) How to measure a turtle. Herpetological Review, 49 (3), 453 - 460.","Smith, M. A. (1931) The Fauna of British India, Including Ceylon and Burma. Volume 1. Loricata and Testudines. Taylor & Francis, London, 185 pp.","Platt, S. G., Lwin, T., Win, M. M., Platt, K., Reh, B., Haislip, N. A. & Rainwater, T. R. (2019) Batagur trivittata (Burmese Roofed Turtle). Sexual size dimorphism. Herpetological Review, 50 (3), 553 - 555.","Moll, E. O. (1986) Survey of the freshwater turtles of India. Part I: The genus Kachuga. Journal of the Bombay Natural History Society, 83 (3), 538 - 552.","Praschag, P., Holloway, R., Georges, A., Packert, M., Hundsdorfer, A. K. & Fritz, U. (2009) A new subspecies of Batagur affinis (Cantor, 1847), one of the world's most critically endangered chelonians (Testudines: Geoemydidae). Zootaxa, 2233 (1), 57 - 68. https: // doi. org / 10.11646 / zootaxa. 2233.1.3","Moll, E. O., Platt, S. G., Chan, E. H., Horne, B. D., Platt, K., Praschag, P., Chen, P. N. & van Dijk, P. P. (2015) Batagur affinis (Cantor 1847) - Southern River Terrapin, Tuntong. Chelonian Research Monograph, 5, 90.1 - 90.17. https: // doi. org / 10.3854 / crm. 5.090. affinis. v 1.2015","Guntoro, J. (2017) TSA Indonesia produces Painted Terrapin hatchlings and new local conservationists. Turtle Survival, 2017, 32 - 33.","Minh, L., McCord, W. P. & Iverson, J. B. (2007) On the paraphyly of the genus Kachuga (Testudinidae: Geoemydidae). Molecular Phylogenetics and Evolution, 45, 398 - 404. https: // doi. org / 10.1016 / j. ympev. 2007.05.002","Praschag, P., Hundsdorfer, A. K. & Fritz, U. (2007) Phylogeny and taxonomy of endangered South and South-east Asian freshwater turtles elucidated by mtDNA sequence variation (Testudines: Geoemydidae: Batagur, Callagur, Hardella, Kachuga, Pangshura). Zoologica Scripta, 36 (5), 429 - 442. https: // doi. org / 10.1111 / j. 1463 - 6409.2007.00293. x","Platt, S. G., Lwin, T., Win, M. M., Platt, K. & Rainwater, T. R. (2017 b) Batagur trivittata (Burmese Roofed Turtle). Description and phenology of sexual dichromatism. Herpetological Review, 48 (3), 616 - 618.","Inbar, M. & Lev-Yadun, S. (2005) Conspicuous and aposematic spines in the animal kingdom. Naturwissenschaften, 92 (1), 170 - 172. https: // doi. org / 10.1007 / s 00114 - 005 - 0608 - 2"]}

The Burmese Roofed Turtle (Batagur trivittata Duméril & Bibron, 1835) is a large (straight-line carapace length [CL] to 620 mm; Platt et al., 2019), aquatic, herbivorous turtle endemic to the major river systems of Myanmar (Smith 1931; TTWG 2017). Although historically widespread and apparently abundant, long-term population declines resulted from chronic egg collecting, subsistence harvesting of adults, and loss of critical nesting habitat (Platt et al. 2017a). By the late 1990s B. trivittata was considered a candidate for Extinct status (Bhupathy et al. 2000) until a living specimen purchased in a Chinese wildlife market came into the possession of an American turtle collector in the early 2000s (Platt et al. 2005; W.P. McCord, pers. comm.). Shortly thereafter, field surveys “rediscovered” two remnant populations in the Dokhtawady and upper Chindwin Rivers (Platt et al. 2005; Kuchling et al. 2006). Intense ex- and in-situ recovery efforts were launched shortly thereafter and continue today (Kuchling & Tint Lwin 2004; Çilingir et al. 2017).

Neogene (Siwalik-aged) deposits from India and Pakistan have yielded many vertebrate fossils, of which most were named during the 19th century, including numerous geoemydid turtles. In contrast to many other faunal components from the Siwaliks, geoemydids have not undergone taxonomic revision for more than a century and most fossils have therefore been believed to correspond to recent taxa. In this study, we conduct a taxonomic revision of all previously described geoemydid material from the Siwalik-age. We propose that all specimens of ‘Clemmys’ from the Siwaliks of Punjab, Pakistan should be identified as Melanochelys sivalensis comb. nov.; that Melanochelys tricarinata var. sivalensis represents a valid species, for which we propose the replacement name Melanochelys tapani to avoid homonymy; that specimens originally identified as Batagur cautleyi and Pangshura flaviventer cannot be identified beyond the generic level; and that many fragmentary palatochelydians cannot be identified to any particular species or genus due to the lack of preserved diagnostic osteological characters. With a few exceptions, the Siwalik fauna mostly corresponds in its distribution to that of the recent fauna, indicating a certain amount of geographic stasis. However, as the stratigraphic provenance of most material is poor, it is not possible to discern meaningful temporal patterns.

Melanochelys tapani nom. nov. Fig. 32 Nicoria tricarinata var. sivalensis Lydekker, 1889b: 100. Geoemyda tricarinata Smith, 1931: 95. Melanochelys tricarinata TEWG, 2015: e.46. Type BMNH 39839, a near complete fossil shell (Fig. 32). Differential osteological diagnosis using shell characters Melanochelys tapani can be differentiated from other Melanochelys species by the following combination of characters: presence of three carapacial keels, a highly domed shell, hexagonal second to fourth vertebrals that are wider than long, and a large cervical scute. Etymology The specific name is in honour of the late Prof. Tapan Roy Chowdhury of the Indian Statistical Institute, the distinguished teacher and researcher of Indian fossils, who established a school of vertebrate palaeontologists in India. Material examined Holotype INDIA ��� Siwalik Hills; Miocene���Pliocene; BMNH 39839. Type locality and horizon Miocene/Pliocene of the Siwalik Hills, likely of India. Description of type BMNH 39839 (Fig. 32), holotype of Melanochelys tapani nom. nov. and subsequent holotype of Nicoria tricarinata sivalensis ��� This is an almost complete carapace associated with a partial plastron from the Miocene/Pliocene Siwalik Hills, likely of India, originally figured and described by Lydekker (1885a: pl. 21.4; 1889b: fig. 21). Our observation of this specimen overall confirms the observations of Lydekker (1889b), but we also illustrate the plastron and a less idealized carapace that lacks sutures (Fig. 32). A part of the anterior margin of the carapace and all posterior peripherals are missing. Total carapace length is approximately 17 cm. Most sulci are preserved on the carapace, but only very few on the plastron. Three longitudinal carapacial keels are present. The lateral keels are closer to the center of the carapace than to the borders. A cervical scute is present. The first vertebral scute is wider than long and contacts the first marginal scutes. The second to fifth vertebral scutes are about the same size and wider than long. The sulcus between the second pleural and third vertebral is straight. The sulcus between the second and third pleural contacts the fifth marginal scute. Most of the plastral surface is not preserved. The anterior plastron margin straight and lacks a median notch. The gular scutes are longer than wide. The pectoroabdominal sulcus contacts the sixth marginal scute. Comments See Discussion for further details., Published as part of Garbin, Rafaella C., Bandyopadhyay, Saswati & Joyce, Walter G., 2020, A taxonomic revision of geoemydid turtles from Siwalik-age of India and Pakistan, pp. 1-67 in European Journal of Taxonomy 652 on pages 47-48, DOI: 10.5852/ejt.2020.652, http://zenodo.org/record/3860236, {"references":["Lydekker R. 1889 b. Catalogue of the Fossil Reptilia and Amphibia in the British Museum (Natural History). Part III. Chelonia. Order of the Trustees, London.","Smith M. A. 1931. The Fauna of British India, Including Ceylon and Burma, Reptilia and Amphibia. Vol. I - Loricata, Testudines. Taylor and Francis, London.","Lydekker R. 1885 a. Indian Tertiary and post-Tertiary Vertebrata, Part III - Siwalik and Narbada Chelonia. Memoirs of the Geological Survey of India, Palaeontologia Indica 10 (3): 155 - 208."]}

Melanochelys sivalensis (Theobald, 1877) comb. nov. Figs 27���31 Bellia sivalensis Theobald, 1877: 44. Clemmys hydaspica Lydekker, 1885a: 172. Clemmys theobaldi Lydekker, 1885a: 173. Clemmys punjabiensis Lydekker, 1885a: 175. Geoemyda trijuga Smith, 1931: 97 (part). Geoemyda sivalensis Smith, 1931: 89. Melanochelys trijuga indopeninsularis TEWG, 2015: e.46. Bellia theobaldi ��� Lydekker 1889a: 58. Clemmys sivalensis ��� Lydekker 1885a: 171. Bellia sivalensis ��� Lydekker 1889a: 58. Type IM E.88, the anterior half of a fossil shell (Fig. 27). Material examined PAKISTAN ��� 1 specimen, holotype of Bellia sivalensis; Punjab, south of Jhand; Middle to Upper Siwalik group of Potwar Plateau; Miocene���Pliocene; IM E.88 ��� 1 specimen, holotype of Clemmys theobaldi; Punjab, Jhand; Middle to Upper Siwalik group of Potwar Plateau; Miocene���Pliocene; IM E.89 ��� 1 specimen; Punjab, Potwar Plateau; Middle to Upper Siwalik group of Potwar Plateau; Miocene��� Pliocene; IM E.90 ��� 1 specimen, holotype of Clemmys punjabiensis; Punjab; Middle to Upper Siwalik group of Potwar Plateau; Miocene���Pliocene; IM E.92 ��� 1 specimen, holotype of Clemmys hydaspica; Punjab, Jhelum district; Middle to Upper Siwalik group of Potwar Plateau; Miocene���Pliocene; IM E.93. Type locality and horizon Miocene/Pliocene, Middle to Upper Siwaliks of Potwar Plateau, south of Jhand, Punjab, Pakistan (see Comments below). Range Miocene/Pliocene, Middle to Upper Siwaliks of Potwar Plateau, Punjab, Pakistan. Differential osteological diagnosis using shell characters Melanochelys sivalensis can be differentiated from other species of Melanochelys by the lack of longitudinal carapacial keels, the presence of hexagonal second to fourth vertebral scutes with concave posterolateral margins that are broader than long and a denser and more rounded shell. Description of material examined IM E.88 (Fig. 27), holotype of Bellia sivalensis ��� This is the anterior half of a shell from the Miocene/ Pliocene, Middle to Upper Siwaliks of Potwar Plateau, south of Jhand, Punjab, Pakistan (see Comments below), initially figured and described by Lydekker (1885a: pl. 20.1). Our observations of the specimen mostly agree with those of Lydekker, although we document more details in the plastron. Most scutes are clearly visible, but only a few bony sutures are apparent. There are no signs of carapacial keels, but notches on the intervertebral sulci indicate the former presence of keels as a juvenile. This is clearly an adult specimen due to its large size (carapace length greater than 20 cm). Growth annuli marks are present on the anterior marginal scutes. The cervical scute is extremely reduced and clasped between the first marginals. First vertebral scute is longer than wide. The second and third vertebrals have rounded lateral margins. The plastron not well preserved, with few visible sutures. The anterior margin of the plastron is straight and the gular scutes are much longer than wide. IM E.89 (Fig. 28), holotype of Clemmys theobaldi ��� This is the anterior half of a shell from the Miocene/ Pliocene, Middle to Upper Siwaliks of Potwar Plateau, Jhand, Punjab, Pakistan (see Comments below) originally figured by Lydekker (1885a: pl. 20.2). Our illustrations mostly agree with those of Lydekker, but we disagree in the presence of a cervical and document the plastron for the first time. Most scutes are clearly visible, but only a few bones are apparent. It is clearly an adult specimen due to its large size (carapace length greater than 20 cm). There are no signs of carapacial keels or growth annuli marks. The cervical scute is present and as long as wide. The first vertebral scute is wider than long and exhibits an anterolateral constriction. The second and third vertebrals have rounded lateral margins. The anterior margin of the plastron is straight and lacks a median notch. The gular scutes are much longer than wide and completely intersect the entoplastron. The humeropectoral sulcus completely crosses the entoplastron posteriorly. The pectoro-abdominal sulcus does not intersect the hyo-hypoplastral suture. IM E.90 (Fig. 29) ��� This is a nearly complete, previously unfigured shell from the Miocene/Pliocene, Middle to Upper Siwaliks of Potwar Plateau, Punjab, Pakistan. Old ontogenetic age combined with poor preservation of the surface makes it near impossible to discern most scutes and sutures. There are no signs of carapacial keels or growth annuli marks. The cervical scute is extremely reduced and placed between the first marginals. The gular scutes are much longer than wide. IM E.92 (Fig. 30), holotype of Clemmys punjabiensis ��� This specimen was collected from the Miocene/ Pliocene, Middle to Upper Siwaliks of Potwar Plateau, Punjab, Pakistan, and was initially figured and described by Lydekker (1885a: pl. 20.3). Our observations of this specimen overall confirm those of Lydekker, although we document some addition sutures. The specimen represents the anterior half of a carapace and articulated fragments of the plastron, and likely represents an adult specimen due to its larger size (carapace length greater than 15 cm). Scutes are clearly discernable, but sutures are restricted to the periphery of the specimen. No carapacial keels or growth annuli marks are visible. The first vertebral scute is as long as wide and has an anterolateral constriction. The second and third vertebral scutes have semi-sinuous lateral margins. The anterior plastral margin is concave. The gular scute is longer than wide. The pectoro-abdominal sulcus does not intersect the hyo-hyoplastral suture. IM E.93 (Fig. 31), holotype of Clemmys hydaspica ��� This specimen consists of a nearly complete carapace and an articulated partial plastron from the Miocene/Pliocene, Middle to Upper Siwaliks of Potwar Plateau, Jhelum district, Punjab, Pakistan. The figures provided by Lydekker (1885a: pl. 20.4) overall agree with our figure, although we see more details in some areas, but less in others. This is a well-preserved specimen that clearly documents most sulci, but only some sutures. It is likely an adult specimen due to its large size (carapace length greater than 15 cm). The plastron is highly damaged and provides no information and we therefore do not figure it. There are no signs of carapacial keels or growth annuli marks. The first vertebral scute is wider than long and has straight lateral margins. The second and third vertebral scute have convex anterolateral and concave posterolateral sides. The sulcus between the first and second pleural, and the second and third pleural contact the fourth and sixth marginal scutes, respectively. The first neural bone only contacts the second costal on the right side, which is probably an anomaly. Comments Two of five specimens listed in this section originate from ���south of Jhand��� (Lydekker 1885a). We were able to locate three places called J(h)and in the Punjab of Pakistan and India: the large town of J(h)and in Attock District, Pakistan, the village of J(h)and in Chakwal District, Pakistan and the village of J(h) and in Jalandhar District, India. The two locations in Pakistan are surrounded by sedimentary exposures that have yielded fossils of Siwaliks age, while the Indian location is located in a flood plain lacking such exposures. We therefore are highly confident that the type locality is positioned in the Punjab of Pakistan. See Discussion for further details regarding the referral of this material to Melanochelys sivalensis., Published as part of Garbin, Rafaella C., Bandyopadhyay, Saswati & Joyce, Walter G., 2020, A taxonomic revision of geoemydid turtles from Siwalik-age of India and Pakistan, pp. 1-67 in European Journal of Taxonomy 652 on pages 40-47, DOI: 10.5852/ejt.2020.652, http://zenodo.org/record/3860236, {"references":["Theobald W. 1877. Description of a new Emydine from the Upper Tertiaries of the Northern Punjab. Records of the Geological Survey of India 10: 43 - 45.","Lydekker R. 1885 a. Indian Tertiary and post-Tertiary Vertebrata, Part III - Siwalik and Narbada Chelonia. Memoirs of the Geological Survey of India, Palaeontologia Indica 10 (3): 155 - 208.","Smith M. A. 1931. The Fauna of British India, Including Ceylon and Burma, Reptilia and Amphibia. Vol. I - Loricata, Testudines. Taylor and Francis, London.","Lydekker R. 1889 a. Notes on Siwalik and Narbada Chelonia. Records of the Geological Survey of India 22 (1): 56 - 59."]}

Batagur kachuga (Gray, 1831) Figs 12���13 Emys lineata Gray, 1831c: 9. Emys kachuga Gray, 1831b: pl. 74. Batagur ellioti Gray, 1862: 264. Kachuga fusca Gray, 1870: 56 (part). Batagur bakeri Lydekker, 1885a: 190. Clemmys (Clemmys) lineata ��� Fitzinger 1835: 123. Batagur lineata ��� Gray 1856 (���1855���): 35. Batagur lineatus ��� G��nther 1864: 39. Batagur elliotti ��� G��nther 1864: 40. Clemmys ellioti ��� Strauch 1865: 88. Kachuga lineata ��� Gray 1870: 56. Batagur kachuga ��� Theobald 1876: 19. ��� Praschag et al. 2007: 439. Kachuga kachuga ��� Smith 1931: 131. Type Unknown (Iverson 1992). Material examined INDIA ��� 1 specimen, holotype of Batagur bakeri; Siwalik Hills, Yamuna-Ganges River basin; Miocene��� Pliocene; BMNH 39835a ��� 1 specimen; Siwalik Hills; Miocene���Pliocene; BMNH R.891. Type locality ��� India ���, restricted by Smith (1931) to ���N. India ��� (Iverson 1992). Occurrence Miocene/Pliocene���Recent. Differential osteological diagnosis using shell characters Batagur kachuga can be differentiated from other species of Batagur by the presence of an elongated fourth vertebral scute that covers the fourth to eighth neural bones and second and third vertebral scutes with straight lateral margins. Description of material examined BMNH 39835a (Fig. 12), holotype of Batagur bakeri ��� This specimen is from Miocene/Pliocene of the Siwalik Hills of India (Yamuna or Ganges River basins) and was presented to BMNH by General W.E. Baker. It was initially figured and described by Lydekker (1885a: pl. 23.2). Our observations mostly agree with those of Lydekker (1885a), although we note an irregularity on the right side of the neural II/III contact and damage that must have incurred over the course of the last century to the anterior margin of the plastron. This is an almost complete specimen, with a well-preserved shell, and perhaps represents an adult female considering its large size (carapace length greater than 50 cm) compared with extant specimens. Most sulci and sutures are visible on the carapace, but only the sulci are apparent on the plastron. The specimen shows no signs of growth annuli or carapacial keels. The cervical scute is present and broader than long. The vertebral scutes are square shaped and with equidimensional anterior and posterior margins. The first vertebral scute has straight lateral margins, but lacks constrictions. The third vertebral scute is broader than long. All available neurals have anteriorly short sides, with exception of the left side of neural II, which displays an abnormality consisting of a supernumerary bone. The anterior margin of the fourth vertebral runs over neural IV. The bony bridge is well developed. The anterior plastron margin is not preserved anymore, but its original configuration is documented in Lydekker (1885a). The pectoroabdominal sulcus with lateral notches suggests the former presence of longitudinal keels that did not intersected the hyo-hypoplastral suture. The anal portion of the plastron not preserved. BMNH R.891 (Fig. 13) ��� This specimen is from the Miocene/Pliocene of the Siwalik Hills, likely of India, and was presented to the British Museum by P.T. Cautley in 1840. It has not been figured previously. This specimen has an incomplete carapace and an almost complete plastron. The carapace consists only of neurals I���IV, a part of costals I���IV and some peripherals. There are no signs of carapacial keels or growth annuli, indicating that this is probably an adult specimen. Two large fontanelles are present, which suggests that this is perhaps a male individual. Neurals II and III are hexagonal with anteriorly short sides. The second and third vertebral scutes have straight lateral sides. The posterior margins of the first and second pleural scutes are straight and cross over costals II and IV, respectively. The anterior and posterior plastral margins are not preserved. A strong axillary notch is present. The entoplastron is not crossed by the humeropectoral sulcus. The hyo-hypoplastral suture contacts peripheral V and does not overlap the pectoroabdominal sulcus. Inguinal scutes are present that contact the femoral scutes. Comments We confirm the previously established identification of these two specimens as Batagur kachuga, due to their large carapace size (greater than 50 cm), highly domed carapace (for BMNH 39835a), presence of a second vertebral that is as long as wide with straight lateral margins, a third vertebral that is hexagonal with short posterolateral sides (visible in BMNH 39835a), and a large plastron with medially converging humeropectoral sulci. This confirms the synonym of Batagur bakeri with Kachuga lineata, as originally proposed by Boulenger (1889) and supported by Lydekker (1889a), TEWG (2015) and TTWG (2017)., Published as part of Garbin, Rafaella C., Bandyopadhyay, Saswati & Joyce, Walter G., 2020, A taxonomic revision of geoemydid turtles from Siwalik-age of India and Pakistan, pp. 1-67 in European Journal of Taxonomy 652 on pages 16-19, DOI: 10.5852/ejt.2020.652, http://zenodo.org/record/3860236, {"references":["Gray J. E. 1831 b. Illustrations of Indian Zoology, Chiefly Selected from the Collection of Major-General Hardwicke. Vol I, part V. Treuttel, Wurtz, Treuttel jun. and Richter, London. https: // doi. org / 10.5962 / bhl. title. 95127","Gray J. E. 1862. Notice of a new species of Batagur from North-Western India. Proceedings of the Zoological Society of London 1863: 253.","Gray J. E. 1870. Supplement to the Catalogue of Shield Reptiles in the Collection of the British Museum, Part I - Testudinata (Tortoises) with figures of the skulls of 36 genera. Order of the Trustees, London.","Lydekker R. 1885 a. Indian Tertiary and post-Tertiary Vertebrata, Part III - Siwalik and Narbada Chelonia. Memoirs of the Geological Survey of India, Palaeontologia Indica 10 (3): 155 - 208.","Fitzinger L. J. 1835. Entwurf einer systematischen Anordnung der Schildkroten nach den Grundsatzen der naturlichen Methode. Annalen des Wiener Museums der Naturgeschichte 1: 105 - 128.","Gray J. E. 1856 (\" 1855 \"). Catalogue of Shield Reptiles in the Collection of the British Museum, Part I - Testudinata (Tortoises). Order of the Trustees, London. https: // doi. org / 10.5962 / bhl. title. 5491","Gunther A. C. L. G. 1864. The Reptiles of British India. Arment Biological Press, Landsville, PA. https: // doi. org / 10.5962 / bhl. title. 5012","Strauch A. 1865. Die Vertheilung der Schildkroten uber den Erdball - ein Zoogeographischer Versuch. Memoires de l'Academie imperiale des Sciences de St. - Petersbourg, 7 ieme serie 8 (13): 1 - 207. https: // doi. org / 10.5962 / bhl. title. 12477","Theobald W. 1876. Descriptive Catalogue of the Reptiles of British India. Thacker, Spink and Co, Kolkata. https: // doi. org / 10.5962 / bhl. title. 54001","Praschag P., Hundsdorfer A. K. & Fritz U. 2007. Phylogeny and taxonomy of endangered South and South-east Asian freshwater turtles elucidated by mtDNA sequence variaton (Testudines: Geoemydidae: Batagur, Callagur, Hardella, Kachuga, Pangshura). Zoologica Scripta 36: 429 - 442. https: // doi. org / 10.1111 / j. 1463 - 6409.2007.00293. x","Smith M. A. 1931. The Fauna of British India, Including Ceylon and Burma, Reptilia and Amphibia. Vol. I - Loricata, Testudines. Taylor and Francis, London.","Iverson J. B. 1992. A Revised Checklist with Distribution Maps of the Turtles of the World. Earlham College, privately printed, Richmond.","Boulenger G. A. 1889. Catalogue of the Chelonians, Rhynchocephalians, and Crocodiles in the British Museum (Natural History). Trustees of the British Museum (Natural History), London.","Lydekker R. 1889 a. Notes on Siwalik and Narbada Chelonia. Records of the Geological Survey of India 22 (1): 56 - 59."]}

Hardella thurjii (Gray, 1831) Figs 19���21 Emys thurjii Gray, 1831a: 22 (nomen novum). Emys thuryi Gray, 1831c: 8 (nomen oblitum). Emys thuji Gray, 1831b (ex errore): pl. 73. Emys flavonigra Lesson, 1831: 120. Emys thugi Gray, 1832 (ex errore): Directions. Clemmys (Clemmys) thurgii Fitzinger, 1835: 123 (nomen novum). Kachuga oldhami Gray, 1869: 200. Batagur falconeri Lydekker, 1885a: 187 Clemmys watsoni Lydekker, 1886a: 541 Hardella indi Gray, 1870: 58. Geoemyda pilgrimi Prasad & Satsangi, 1967. Emys thurgii ��� Gray 1844: 17. Emys thurgi ��� Gray 1856 (���1855���): 21. Clemmys thurgii ��� Strauch 1862: 32. Batagur thurgii ��� Theobald 1868: 12. Hardella thurgi ��� Gray 1870: 58. Batagur (Hardella) thurgi ��� Anderson 1879: 764. Hardella thurjii ��� Siebenrock 1909: 456. Hardella thurgi ��� Smith 1931: 50. Hardella thurjii thurjii ��� Wermuth & Mertens 1977: 40. Hardella thurjii indi ��� Wermuth & Mertens 1977: 40. Hardella thurji ��� Pritchard 1979 (ex errore): 193. Hardella thurji thurji ��� Obst 1985: 221. Type Unknown (Iverson 1992). Material examined INDIA ��� 1 specimen, holotype of Batagur falconeri; Siwalik Hills; Miocene���Pliocene; BMNH 39835 ��� 1 specimen; Siwalik Hills; Miocene���Pliocene; BMNH R.890 ��� 1 specimen, holotype of Clemmys watsoni; Gulf of Cambay, Gujarat, Piram Island; Late Miocene���Pliocene; BMNH R.748. Type locality ��� India ��� (Iverson 1992). Occurrence Miocene/Pliocene ��� Recent. Differential osteological diagnosis using shell characters Hardella thurjii can be differentiated from other geoemydids by large carapace size (up to 60 cm in females), presence of hexagonal, nearly square second to fourth vertebrals, a first vertebral scute that is wider posteriorly, presence of an inflection at the margin of the gulohumeral sulcus, and an entoplastron that is not intersected by the humeropectoral sulcus. Description of material examined BMNH 39835 (Fig. 19), holotype of Batagur falconeri ��� This is an almost complete shell, exceptionally well preserved, from the Miocene/Pliocene of the Siwalik Hills, likely of India, originally figured and described in three views by Lydekker (1885a: pls 23.1, 24.4). Our figures are overall comparable to those of Lydekker, although we see more details to the peripherals (Fig. 19). The specimen clearly represents an adult female specimen based on overall size (carapace length greater than 40 cm). A small keel is present on neurals IV and VI. There are no signs of growth annuli. All neural bones have short anterior sides. The third neural is half one and a half time longer than the other neural bones. The cervical scute is present, longer than wide. The first vertebral scute is broader posteriorly and narrows anteriorly. The second and third vertebrals are much broader than the other vertebral scutes. The fifth vertebral has an anterolateral constriction. Marginal scutes IV���X overlap the adjacent costal bones. The anterior and posterior plastral margins are not preserved. The pectoroabdominal sulcus has lateral notches, suggesting the former presence of longitudinal keels as a juvenile. The pectoroabdominal sulcus does not intersect the hyo-hypoplastron suture and contacts the fifth marginal scute. Axillary and inguinal scutes are likely present. BMNH R.748 (Fig. 20), holotype of Clemmys watsoni ��� This specimen originates from the Late Miocene���Pliocene of Piram Island, Gulf of Cambay, Gujarat, India, was presented to the BMNH in 1886 by Col. J.W. Watson and figured and described in a small contribution from Lydekker (1886a: pl. 1). Our illustrations in three views overall confirm most of Lydekker���s observations, but we see fewer details along the neural column and the damaged portions of the costals and peripherals. The specimen is almost complete, full size, and misses some lateral peripheral bones (right and left), the anterior plastral margin, the posterior plastral lobe, and the right bridge (Fig. 20). The specimen is likely an adult considering its size (carapace length greater than 15 cm) and perhaps a male specimen, as modern male individuals of H. thurjii reach up to 18 cm and lack intercostal fontanelles (Das & Bhupathy 2009a). Most sulci and sutures of the carapace are visible, as well as a knob on neurals IV and VIII, indicating the presence of a median keel. Growth annuli are present. The cervical scute is as wide as long and lacks a posterior notch. The first vertebral scute is wider than long, its lateral sides converge anteriorly and lack an anterolateral constriction, and it contacts the first marginal scute. The neural bones are hexagonal and anteriorly short-sided. The third vertebral has straight lateral sides and the posterior margin has an anteriorly oriented inflection that crosses the suture between neural bones III���IV. The fifth vertebral scute has an anterolateral constriction. The pygal bone is completely intersected by the twelfth intermarginal sulcus. The humeropectoral sulcus is located posterior to the entoplastron. The pectoral scute contacts the fifth marginal. For a more extensive description of this specimen, we refer to Lydekker (1886a). BMNH R.890 (Fig. 21) ��� This specimen is from the Miocene/Pliocene of the Siwalik Hills, likely of India, was purchased by P.T. Cautley in 1840, but remained unfigured to date. It is a crushed, partial specimen, that probably represents an adult female considering its large size (carapace length greater than 20 cm). Both the anterior and posterior margins of the carapace are missing. Neural bones III and IV have a knob, indicating the presence of a median carapacial keel. Neural bones II���IV are hexagonal with anterior short sides. The first vertebral scute has anteriorly converging lateral sides. The third vertebral scute has straight lateral sulci. The posterior margins of the first and second pleurals are straight, placed over costals II and IV, respectively, and lack an anterior projection. Both the anterior and posterior plastral margins are not preserved. The plastron has two lateral longitudinal keels, which cross the lateral sides of the hyo- and hyoplastra. The humeropectoral sulcus is apparently located posterior to the entoplastron, which is not preserved. Inguinal scutes are likely present that contact the femoral scute. Comments We here attribute these specimens to Hardella thurjii based on the presence of a short median keel, large and square second and third vertebrals that are about the same width as the fourth vertebral scute, large inguinal scute, and an entoplastron that is not intersected by the humeropectoral sulcus. This confirms the synonymy of B. falconeri and C. watsoni with H. thurjii, as first suggested by Boulenger (1889) and later supported by Lydekker (1889a), TEWG (2015) and TTWG (2017). We furthermore follow Das (1994) by recognizing the synonymy of Geoemyda pilgrimi with H. thurjii (followed by TEWG 2015 and TTWG 2017) although we did not study the holotype of this species firsthand., Published as part of Garbin, Rafaella C., Bandyopadhyay, Saswati & Joyce, Walter G., 2020, A taxonomic revision of geoemydid turtles from Siwalik-age of India and Pakistan, pp. 1-67 in European Journal of Taxonomy 652 on pages 28-33, DOI: 10.5852/ejt.2020.652, http://zenodo.org/record/3860236, {"references":["Gray J. E. 1831 a. Synopsys Reptilium: or Short Descriptions of the Species of the Reptiles. Part I - Cataphracta. Tortoises, Crocodiles, and Enaliosaurus. Treuttel, Wurz and Co, London.","Gray J. E. 1831 b. Illustrations of Indian Zoology, Chiefly Selected from the Collection of Major-General Hardwicke. Vol I, part V. Treuttel, Wurtz, Treuttel jun. and Richter, London. https: // doi. org / 10.5962 / bhl. title. 95127","Lesson R. P. 1831. Catalogue des reptiles qui font partie d`une collection zoologique recueillie dans l`Inde continentale ou en Afrique, et apportee en Franc par M. Lamare-Piquot. Bulletin des Sciences naturelles et Geologie de Paris 25: 119 - 123.","Gray J. E. 1832. Illustrations of Indian Zoology, Chiefly Selected from the Collection of Major-General Hardwicke. Vol II, part XI. Adolphus Richter and Co., London. https: // doi. org / 10.5962 / bhl. title. 95127","Fitzinger L. J. 1835. Entwurf einer systematischen Anordnung der Schildkroten nach den Grundsatzen der naturlichen Methode. Annalen des Wiener Museums der Naturgeschichte 1: 105 - 128.","Gray J. E. 1869. Notes on the families and genera of tortoises (Testudinata), and on the characters afforded by the study of their skulls. Proceedings of the Zoological Society of London 1869: 165 - 225. https: // doi. org / 10.1111 / j. 1469 - 7998.1869. tb 07312. x","Lydekker R. 1885 a. Indian Tertiary and post-Tertiary Vertebrata, Part III - Siwalik and Narbada Chelonia. Memoirs of the Geological Survey of India, Palaeontologia Indica 10 (3): 155 - 208.","Lydekker R. 1886 a. On a new Emydine chelonian from the Pliocene of India. Quarterly Journal of the Geological Society 42: 540 - 541. https: // doi. org / 10.1144 / gsl. jgs. 1886.042.01 - 04.54","Gray J. E. 1870. Supplement to the Catalogue of Shield Reptiles in the Collection of the British Museum, Part I - Testudinata (Tortoises) with figures of the skulls of 36 genera. Order of the Trustees, London.","Prasad K. N. & Satsangi P. P. 1967. On a new fossil chelonian from the Siwalik beds of Himachal Pradesh. Records of the Geological Survey of India 95: 533 - 536.","Gray J. E. 1844. Catalogue of the Tortoises, Crocodiles and Amphisbaenians, in the Collection of the British Museum. Order of the Trustees, London. https: // doi. org / 10.5962 / bhl. title. 21639","Gray J. E. 1856 (\" 1855 \"). Catalogue of Shield Reptiles in the Collection of the British Museum, Part I - Testudinata (Tortoises). Order of the Trustees, London. https: // doi. org / 10.5962 / bhl. title. 5491","Strauch A. 1862. Chelonologische Studien, mit besonderer Beziehung auf die Schildkrotensammlung der Kaiserlichen Akademie der Wissenschaften zu St. - Petersburg. Memoires de l'Academie imperiale des Sciences de St. - Petersbourg, 7 ieme serie 5 (7): 1 - 197.","Theobald W. 1868. Catalogue of reptiles in the Museum of the Asiatic Society of Bengal. Journal of the Asiatic Society of Bengal, Extra Number. Baptist Mission Press, Kolkata. https: // doi. org / 10.5962 / bhl. title. 5477","Anderson J. A. 1879. Anatomical and Zoological Researches: Comprising an Account of the Zoological Results of the two Expeditions to Western Yunnan in 1868 and 1875; and a Monograph of the two Cetacean Genera, Platanista and Orcella. Bernard Quaritch, London.","Siebenrock F. 1909. Synopsis der rezenten Schildkroten, mit Berucksichtigung der in historischer Zeit ausgestorbenen Arten. Zoologische Jahrbucher Suppl. 10 (3): 427 - 618. https: // doi. org / 10.5962 / bhl. title. 12485","Smith M. A. 1931. The Fauna of British India, Including Ceylon and Burma, Reptilia and Amphibia. Vol. I - Loricata, Testudines. Taylor and Francis, London.","Wermuth H. & Mertens R. 1977. Testudines, Crocodylia, Rhynchocephalia. Das Tierreich. Walter de Gruyter, Berlin.","Pritchard P. C. H. 1979. Encyclopedia of Turtles. T. F. H. Publications, New Jersey.","Iverson J. B. 1992. A Revised Checklist with Distribution Maps of the Turtles of the World. Earlham College, privately printed, Richmond.","Das I. & Bhupathy S. 2009 a. Hardella thurjii (Gray 1831) - Crowned River Turtle. Chelonian Research Monographs 5: 023.1 - 023.6. https: // doi. org / 10.3854 / crm. 5.023. thurjii. v 1.2009","Lydekker R. 1889 b. Catalogue of the Fossil Reptilia and Amphibia in the British Museum (Natural History). Part III. Chelonia. Order of the Trustees, London.","Boulenger G. A. 1889. Catalogue of the Chelonians, Rhynchocephalians, and Crocodiles in the British Museum (Natural History). Trustees of the British Museum (Natural History), London.","Lydekker R. 1889 a. Notes on Siwalik and Narbada Chelonia. Records of the Geological Survey of India 22 (1): 56 - 59.","Das I. 1994. The identity of the Plio-Pleistocene turtle, Geoemyda pilgrimi Prasad and Satsangi, 1967 (Testudines: Cryptodira: Bataguridae). Hamadryad 19: 41 - 46."]}

Genus Batagur Gray, 1856 Figs 10���11 Type species Batagur baska (Gray, 1830). Differential osteological diagnosis using shell characters A representative of Batagur can be diagnosed by the presence of a large carapace size (median carapace length of more than 40 cm in adults), a well-developed bridge, well-developed axillary and inguinal buttresses, neural scutes with anterior short sides, a long third neural bone, a short, anteriorly truncated gular scute, an entoplastron that is not intersected by the humeropectoral sulcus and a short anal notch. Material examined INDIA ��� 1 specimen, holotype of Batagur cautleyi Lydekker, 1885; Siwalik Hills; Miocene���Pliocene; BMNH 39834. COUNTRY UNKNOWN ��� 1 specimen; IM E.176. Description BMNH 39834 (Fig. 10), holotype of Batagur cautleyi ��� This an almost complete shell from the Miocene/ Pliocene of the Siwalik Hills, likely of India, showing the majority of sulci and of the sutures along the center of the carapace. It was originally figured by Lydekker (1885a: pl. 24.1), but our observations reveal many additional sutures. There are no signs of growth annuli. The specimen likely is an adult female due to its large size (carapace length greater than 54 cm). A small protuberance on neural IV suggests the previous presence of a median keel. A cervical scute is present. The first vertebral scute is longer than wide and shows a lateral constriction. Neurals II���IV have anteriorly short sides, but neural V has a short left posterior side, which is probably abnormal. The sulcus between the second pleural and the third vertebral is almost straight. The anterior plastral margin is straight and lacks an inflection on the contact of the gularohumeral sulcus. The pectoroabdominal sulcus contacts the sulcus between fifth and sixth marginal scutes. The anal notch is not preserved in this specimen. No plastral sutures can be seen. IM E.176 (Fig. 11) ��� This is a well-preserved specimen that lacks provenance data and appears to be unpublished. The majority of sutures and sulci are clearly visible. This is an adult specimen due to its large size (carapace length greater than 35 cm), but its gender is unknown. A median carapacial keel is present on the posterior parts of the carapace and is strongly marked over neurals IV and VI. The carapace is rather domed at its center, with no signs of growth annuli. Vertebral scutes are subrectangular and have equally sized anterior and posterior margins. Neurals II to VI are anteriorly shortsided. Gular scutes appear to be wider than long and overlap part of the entoplastron. The humeropectoral sulcus is located posterior to the entoplastron. The pectoroabdominal sulcus does not intersect the hyohypoplastron suture and has two lateral notches, suggesting the former presence of parasagital plastral keels at the bridge. Comments We attributed BMNH 39834 and IM E.176, the type series of Batagur cautleyi, to Batagur indet. based on their massive size, highly domed carapace, and presence of an axillary notch and rather straight anterior plastral margin. However, we cannot identify these specimens to species level even though they display unique character combinations. See Discussion (Batagur cautleyi from the Siwalik Hills) for additional details., Published as part of Garbin, Rafaella C., Bandyopadhyay, Saswati & Joyce, Walter G., 2020, A taxonomic revision of geoemydid turtles from Siwalik-age of India and Pakistan, pp. 1-67 in European Journal of Taxonomy 652 on pages 13-16, DOI: 10.5852/ejt.2020.652, http://zenodo.org/record/3860236, {"references":["Gray J. E. 1856 (\" 1855 \"). Catalogue of Shield Reptiles in the Collection of the British Museum, Part I - Testudinata (Tortoises). Order of the Trustees, London. https: // doi. org / 10.5962 / bhl. title. 5491","Gray J. E. 1830. A synopsis of the species of class Reptilia. In: Griffith E. & Pidgeon E. (eds) The Class Reptilia Arranged by the Baron Cuvier, with Specific Descriptions. In: Griffith E. (ed.) The Animal Kingdom Arranged in Conformity with its Organization by the Baron Cuvier, with Additional Descriptions of all the Species Hitherto Named, and of Many not Before noticed. Vol. 9. Reptilia. Whittaker, Treacher and Co, London.","Lydekker R. 1885 a. Indian Tertiary and post-Tertiary Vertebrata, Part III - Siwalik and Narbada Chelonia. Memoirs of the Geological Survey of India, Palaeontologia Indica 10 (3): 155 - 208."]}

Genus Pangshura Gray, 1856 Figs 16���18 Type species Pangshura tecta (Gray, 1830). Differential osteological diagnosis using shell characters A member of Pangshura can be diagnosed by the presence of a tectiform carapace, strong median carapacial keel, a pleural I/II sulcus with an anteromedial process, a pleural IV/vertebral V sulcus contacting the tenth marginal, a fourth vertebral scute that is much longer than wide and that is strongly constricted anteriorly, and an octagonal fourth neural (Garbin et al. 2018). Material examined INDIA ��� 1 specimen; Yamuna-Ganges River basin, Siwalik Hills; Miocene���Pliocene; BMNH 39837 ��� 1 incomplete specimen; Siwalik Hills; Miocene���Pliocene; BMNH 17435 ��� 1 specimen, original of ��� Emys namadicus ���; Madhya Pradesh, Central Narmada Valley, Muwar and Doomar Villages; Late Pleistocene; IM E.110. Description of material examined BMNH 39837 (Fig. 16) ��� This is an almost complete shell from the Miocene/Pliocene of the Yamuna or Ganges River basins in the Siwalik Hills of India originally figured and described in dorsal view by Lydekker (1885a: pl. 22.3). The specimen is not particularly well preserved, but many more sutures are visible than apparent from the figure of Lydekker. The size (carapace length greater than 15 cm) indicates that it represents an adult female. A small keel is present between neural II and suprapygal II. There are no signs of growth annuli. Neural bones II, III, and VII are hexagonal with anterior short sides. Neural IV is octagonal. The third vertebral scute has a strong posterior keel, but no midline prominence. The posterior margin of the first and second pleural scutes have long finger-like anterior projections that intersect costal I and III, respectively. The fifth marginal scute contacts costal bones III and IV. The anterior and posterior plastral margins are not completely preserved. The entoplastron is not intersected by the humeropectoral sulcus. The pectoroabdominal sulcus has lateral notches, which suggest the former presence of longitudinal keels, does not intersect the hyo-hypoplastral suture, and contacts the fifth marginal scute. Axillary and inguinal scutes were likely present. The anal notch is deep. BMNH 17435 (Fig. 17) ��� This is a small incomplete specimen (total length of 9 cm) from the Miocene/ Pliocene of the Siwalik Hills, likely of India, that was presented to BMNH by P.T. Cautley in 1840 and figured and described in dorsal view by Lydekker (1885a: pl. 22.1). The specimen probably represents an adult male based on its small size and consists of the center of the shell, missing the anterior, posterior as well as part of lateral carapace margins. Some additional sutures are apparent relative to the original figure of Lydekker (1885a). A strong median keel is present, running from the most anterior to the most posterior region of the specimen. The carapace has a tectiform shape in anterior view (not illustrated). There are no signs of growth annuli or intercostal fontanelles. A cervical scute is present. The first vertebral scute is small, with straight lateral margins. The second vertebral scute is hexagonal, with shorter posterolateral margins, and a straight sulcus between the second and third vertebral. The third vertebral scute is pentagonal, has straight lateral margins and a small posterior projection into the fourth vertebral. The fourth vertebral scute is only partially preserved, but constricted anteriorly as strongly as other representatives of Pangshura. The interpleural sulcus I���II lies over the suture between costal I and II, and intersects it anteriorly, without a finger-like projection. The interpleural sulcus II���III has a small anterior projection that almost intersects the suture between costal III and IV. The anterior and posterior plastral margins are not preserved. The entoplastron is not intersected by the humeropectoral sulcus. The hyo-hypoplastral suture does not overlap the pectoroabdominal sulcus and contacts peripheral V laterally. The axillary and inguinal scutes are likely present. The fourth, fifth, and sixth marginals form the well-developed bridge. IM E.110 (Fig. 18), original of ��� Emys namadicus ��� Theobald, 1860 (nomen nudum) ��� This specimen is a well-preserved shell of a small individual (total length of 8 cm) from the latest Pleistocene of Muwar-Doomar, Central Narmada Valley, India, originally named by Theobald (1860) but only later figured in dorsal view and described by Lydekker (1885a: pl. 22.2). Although we observe fewer sulci and sutures than originally documented by Lydekker, most are well preserved. The presence of a hyo- hypoplastral fontanelle suggests that this is likely a juvenile specimen. A strong median carapacial keel is present, which is at its highest at the posterior region of the third vertebral scute. There are no signs of growth annuli marks. The first vertebral scute has a wide anterior margin followed by a constriction of the lateral margins. The second vertebral scute is hexagonal, subquadrangular, and as long as broad. The third vertebral scute is pentagonal and pointed posteriorly. The fourth vertebral scute is rhomboid, with a slight anterior constriction, and its posterior margin overlaps the suture between the eighth neural bone and the first suprapygal. The sulcus between the first and second pleural scutes is almost straight, overlapping the suture between costal I and II. The sulcus between the second and third pleurals is directed anteriorly in the dorsal portion, without an anterior projection or finger-like process. The anterior and posterior plastral margins are missing. The bridge is well developed with the fourth, fifth, and sixth marginals overlapping the hyo- and hypoplastra. The pectoroabdominal sulcus does not intersect the hyo-hypoplastral suture. The inguinal scute is likely present. Comments The three herein referred specimens in our opinion lack characters that would allow identifying them to species level. This contradicts in part the original assessments of Lydekker (1885a). See Discussion (Pangshura specimens section) for further details., Published as part of Garbin, Rafaella C., Bandyopadhyay, Saswati & Joyce, Walter G., 2020, A taxonomic revision of geoemydid turtles from Siwalik-age of India and Pakistan, pp. 1-67 in European Journal of Taxonomy 652 on pages 23-27, DOI: 10.5852/ejt.2020.652, http://zenodo.org/record/3860236, {"references":["Gray J. E. 1856 (\" 1855 \"). Catalogue of Shield Reptiles in the Collection of the British Museum, Part I - Testudinata (Tortoises). Order of the Trustees, London. https: // doi. org / 10.5962 / bhl. title. 5491","Gray J. E. 1830. A synopsis of the species of class Reptilia. In: Griffith E. & Pidgeon E. (eds) The Class Reptilia Arranged by the Baron Cuvier, with Specific Descriptions. In: Griffith E. (ed.) The Animal Kingdom Arranged in Conformity with its Organization by the Baron Cuvier, with Additional Descriptions of all the Species Hitherto Named, and of Many not Before noticed. Vol. 9. Reptilia. Whittaker, Treacher and Co, London.","Garbin R. C., Ascarrunz E. & Joyce W. G. 2018. Polymorphic characters in the reconstruction of the phylogeny of geoemydid turtles. Zoological Journal of the Linnean Society 184: 896 - 918. https: // doi. org / 10.1093 / zoolinnean / zlx 106","Lydekker R. 1885 a. Indian Tertiary and post-Tertiary Vertebrata, Part III - Siwalik and Narbada Chelonia. Memoirs of the Geological Survey of India, Palaeontologia Indica 10 (3): 155 - 208.","Gunther A. C. L. G. 1864. The Reptiles of British India. Arment Biological Press, Landsville, PA. https: // doi. org / 10.5962 / bhl. title. 5012","Theobald W. 1860. On the tertiary and alluvial deposits of the central portion of the Nerbudda valley. Memoirs of the Geological Survey of India 2: 279 - 298."]}

Genus Melanochelys Gray, 1869 Type species Melanochelys trijuga (Schweigger, 1812). Differential osteological diagnosis using shell characters A member of Melanochelys can be diagnosed by having a small to medium sized, oval carapace (up to 30 cm in length), presence of three longitudinal keels, an octagonal second neural, neural III���VI with posterior short sides, a large first vertebral with lateral constriction, hexagonal second to fourth vertebral scutes, gular scutes that are longer than wide and an entoplastron that is intersected by the humeropectoral sulcus., Published as part of Garbin, Rafaella C., Bandyopadhyay, Saswati & Joyce, Walter G., 2020, A taxonomic revision of geoemydid turtles from Siwalik-age of India and Pakistan, pp. 1-67 in European Journal of Taxonomy 652 on page 40, DOI: 10.5852/ejt.2020.652, http://zenodo.org/record/3860236, {"references":["Gray J. E. 1869. Notes on the families and genera of tortoises (Testudinata), and on the characters afforded by the study of their skulls. Proceedings of the Zoological Society of London 1869: 165 - 225. https: // doi. org / 10.1111 / j. 1469 - 7998.1869. tb 07312. x"]}

Genus Geoclemys Gray, 1856 Type species Geoclemys hamiltonii (Gray, 1830). Differential osteological diagnosis using shell characters See Geoclemys hamiltonii below., Published as part of Garbin, Rafaella C., Bandyopadhyay, Saswati & Joyce, Walter G., 2020, A taxonomic revision of geoemydid turtles from Siwalik-age of India and Pakistan, pp. 1-67 in European Journal of Taxonomy 652 on page 33, DOI: 10.5852/ejt.2020.652, http://zenodo.org/record/3860236, {"references":["Gray J. E. 1856 (\" 1855 \"). Catalogue of Shield Reptiles in the Collection of the British Museum, Part I - Testudinata (Tortoises). Order of the Trustees, London. https: // doi. org / 10.5962 / bhl. title. 5491","Gray J. E. 1830. A synopsis of the species of class Reptilia. In: Griffith E. & Pidgeon E. (eds) The Class Reptilia Arranged by the Baron Cuvier, with Specific Descriptions. In: Griffith E. (ed.) The Animal Kingdom Arranged in Conformity with its Organization by the Baron Cuvier, with Additional Descriptions of all the Species Hitherto Named, and of Many not Before noticed. Vol. 9. Reptilia. Whittaker, Treacher and Co, London."]}

Pangshura tatrotia Joyce & Lyson, 2010 Type YPM 4127, a near complete fossil shell. Material examined Holotype PAKISTAN ��� Punjab, Potwar Plateau, 2 miles north-east of Padhri; Tatrot Formation; Late Pliocene; YPM 4127. Type locality Yale North India Expedition locality 99, about two miles north-east of Padhri, Potwar Plateau, Punjab, Pakistan. Occurrence Tatrot Formation, Late Pliocene (3.6 to 2.6 Ma, possibly Plio���Pleistocene boundary). Differential osteological diagnosis using shell characters Pangshura tatrotia can be differentiated from other species of Pangshura by the presence of a strong median keel projection on both the second and third vertebral, and a first vertebral scute that is constricted anteriorly. Description of the type YPM 4127, holotype of Pangshura tatrotia ��� This is an almost complete, well-preserved specimen, with a tectiform carapace from the Early Pliocene Tatrot Formation of the Potwar Plateau of Punjab, Pakistan. The posterior peripheral bones and left peripheral bones IV���VI are missing. A median, wellpronounced carapacial keel is present from neurals II to VIII. Neural bones II, III, VI���VIII are hexagonal with anterior short sides. Neural IV is octagonal, with both anterior and posterior sides short. Neural V is quadrangular, without short sides. The fourth vertebral scute runs from neural IV to VIII and has a strong anterior bottle-neck-shaped constriction. The posterior margins of the first and second pleural scutes run over costals II and IV, respectively, and have a strong anterior projection that crosses to the anterior costal bone. The pygal bone is completely divided by the twelfth intermarginal sulcus. Parts of the anterior and posterior plastral margins are missing. The entoplastron is intersected anteriorly by the gularohumeral sulcus, but not by the humeropectoral sulcus. The hyo-hypoplastral suture contacts peripheral V and does not overlap with the pectoroabdominal sulcus. The fifth and sixth marginal scutes form the bridge and overlap onto the hyoplastra. Only the sixth marginal scute overlaps with the hypoplastron. A large inguinal scute is present, likely contacting the femoral scute. For a more comprehensive description of this specimen, refer to Joyce & Lyson (2010). Comments Pangshura tatrotia was only recently named based on a well-preserved shell that documents with confidence a morphotype different from all extant representatives of Pangshura. The specimen is furthermore associated with quality locality data. We therefore here find the validity of this species to be unproblematic., Published as part of Garbin, Rafaella C., Bandyopadhyay, Saswati & Joyce, Walter G., 2020, A taxonomic revision of geoemydid turtles from Siwalik-age of India and Pakistan, pp. 1-67 in European Journal of Taxonomy 652 on pages 27-28, DOI: 10.5852/ejt.2020.652, http://zenodo.org/record/3860236, {"references":["Joyce W. G. & Lyson T. R. 2010. Pangshura tatrotia, a new species of pond turtle (Testudinoidea) from the Pliocene Siwaliks of Pakistan. Journal of Systematic Palaeontology 8 (3): 449 - 458. https: // doi. org / 10.1080 / 14772019.2010.500879"]}

Batagur dhongoka (Gray, 1832) Figs 14���15 Emys dhongoka Gray, 1832: pl. 60. Emys duvaucelli Dum��ril & Bibron, 1835: 334. Kachuga hardwickii Gray, 1869: 202. Batagur durandi Lydekker, 1885a: 192. Batagur dhongoka ��� Gray 1855 (1856): 36. ��� Praschag et al. 2007: 439. Clemmys dhongoka ��� Strauch 1862: 33. Dhongoka hardwickii ��� Gray 1870: 56. Batagur duvaucelli ��� Anderson 1879: 738. Kachuga dhongoka ��� Boulenger 1889: 56. Type Unknown (Iverson 1992). Material examined INDIA ��� 1 specimen, holotype of Batagur durandi; Siwalik Hills; Miocene���Pliocene; BMNH 39841. COUNTRY UNKNOWN ��� 1 specimen; IM W19/173. Type locality Not stated originally, restricted by Smith (1931) to ���N. India ��� (Iverson 1992). Occurrence Miocene/Pliocene ��� Recent. Differential osteological diagnosis using shell characters Batagur dhongoka can be differentiated from other Batagur species by the presence of an elongated fourth vertebral scute that overlaps four neural bones, a second vertebral scute with a posterior protrusion into the third vertebral, a straight humeropectoral sulcus, and a gulohumeral sulcus that forms a right angle. Description of material examined BMNH 39841 (Fig. 14), holotype of Batagur durandi ��� This is an almost complete, well-preserved specimen from Miocene/Pliocene of the Siwalik Hills, probably of India, that was presented to BMNH by P.T. Cautley. The original figure by Lydekker (1885a: pl. 24.2) overall compares well to our observations, but we note differences in the shape of vertebrals III and IV and the presence of peripherals, and we document the plastron for the first time (Fig. 14 C���D).A portion of the anterior margin of the carapace and some posterior left peripheral bones are missing. The specimen clearly represents an adult female due to its large size (carapace length greater than 40 cm). A median longitudinal carapacial keel is present, which is elevated in the posterior region of the second vertebral scute. All neural bones are hexagonal and anteriorly short-sided. The first to fourth neural bones are about the same size and longer than wide. The fifth to eighth neural bones are wider than long. The seventh neural is anomalously divided into two elements. The first vertebral scute is bell-shaped and has a small anterolateral constriction. The second vertebral scute has a deep protrusion along its posterior margin into the third vertebral. The third vertebral scute has a smaller protrusion into the fourth vertebral scute. The fourth vertebral is twice as long than wide and its anterior margin intersects the fourth neural. The sulcus between the first and second pleural forms a deep anterolateral projection onto the first costal bone. The fifth and sixth marginal scutes overlap part of the costal bones. The anterior plastral margin is not completely preserved. The entoplastron is not intersected by the humeropectoral sulcus. the pectoroabdominal sulcus contacts the fifth marginal scute on one side of the specimen, but the sixth marginal on the other. Both the fifth and sixth marginal scutes overlap the hyoplastron. The hyo-hypoplastral suture and pectoroabdominal sulcus do not overlap or coincide. The xiphiplastra have a small, rounded anal notch. IM W19/173 (Fig. 15) ��� This is a well-preserved specimen that lacks provenance data and that appears to be unpublished. The majority of sutures and sulci of the carapace are clearly visible. The specimen likely represents an adult female due to its large size (carapace length greater than 40 cm). The carapace is highly domed at its center. A median carapacial keel is present, with protrusions at the posterior margins of the second and third vertebral scutes. The neural bones, likely the sixth to eighth, are anteriorly short-sided. The sulcus between the first and the second pleurals and the second and third pleurals are positioned over the second and fourth costal bones, respectively. The plastron is damaged and thus not shown here. Comments Here, we attribute these two specimens to Batagur dhongoka based on the protrusion of the second vertebral into the third, a medially short third vertebral scute, a fourth vertebral scute that is much longer than wide, and a large plastron with straight humeropectoral sulci that do not cross the entoplastron (noticeable on BMNH 39841). This confirms the synonym of B. durandi with Batagur dhongoka, as first suggested by Boulenger (1889) and later supported by Lydekker (1889a), TEWG (2015) and TTWG (2017)., Published as part of Garbin, Rafaella C., Bandyopadhyay, Saswati & Joyce, Walter G., 2020, A taxonomic revision of geoemydid turtles from Siwalik-age of India and Pakistan, pp. 1-67 in European Journal of Taxonomy 652 on pages 20-22, DOI: 10.5852/ejt.2020.652, http://zenodo.org/record/3860236, {"references":["Gray J. E. 1832. Illustrations of Indian Zoology, Chiefly Selected from the Collection of Major-General Hardwicke. Vol II, part XI. Adolphus Richter and Co., London. https: // doi. org / 10.5962 / bhl. title. 95127","Dumeril A. M. C. & Bibron G. 1835. Erpetologie generale ou Histoire naturelle complete des Reptiles - II. Cheloniens. Librairie encyclopedique de Roret, Paris.","Gray J. E. 1869. Notes on the families and genera of tortoises (Testudinata), and on the characters afforded by the study of their skulls. Proceedings of the Zoological Society of London 1869: 165 - 225. https: // doi. org / 10.1111 / j. 1469 - 7998.1869. tb 07312. x","Lydekker R. 1885 a. Indian Tertiary and post-Tertiary Vertebrata, Part III - Siwalik and Narbada Chelonia. Memoirs of the Geological Survey of India, Palaeontologia Indica 10 (3): 155 - 208.","Praschag P., Hundsdorfer A. K. & Fritz U. 2007. Phylogeny and taxonomy of endangered South and South-east Asian freshwater turtles elucidated by mtDNA sequence variaton (Testudines: Geoemydidae: Batagur, Callagur, Hardella, Kachuga, Pangshura). Zoologica Scripta 36: 429 - 442. https: // doi. org / 10.1111 / j. 1463 - 6409.2007.00293. x","Strauch A. 1862. Chelonologische Studien, mit besonderer Beziehung auf die Schildkrotensammlung der Kaiserlichen Akademie der Wissenschaften zu St. - Petersburg. Memoires de l'Academie imperiale des Sciences de St. - Petersbourg, 7 ieme serie 5 (7): 1 - 197.","Gray J. E. 1870. Supplement to the Catalogue of Shield Reptiles in the Collection of the British Museum, Part I - Testudinata (Tortoises) with figures of the skulls of 36 genera. Order of the Trustees, London.","Anderson J. A. 1879. Anatomical and Zoological Researches: Comprising an Account of the Zoological Results of the two Expeditions to Western Yunnan in 1868 and 1875; and a Monograph of the two Cetacean Genera, Platanista and Orcella. Bernard Quaritch, London.","Boulenger G. A. 1889. Catalogue of the Chelonians, Rhynchocephalians, and Crocodiles in the British Museum (Natural History). Trustees of the British Museum (Natural History), London.","Iverson J. B. 1992. A Revised Checklist with Distribution Maps of the Turtles of the World. Earlham College, privately printed, Richmond.","Smith M. A. 1931. The Fauna of British India, Including Ceylon and Burma, Reptilia and Amphibia. Vol. I - Loricata, Testudines. Taylor and Francis, London.","Lydekker R. 1889 a. Notes on Siwalik and Narbada Chelonia. Records of the Geological Survey of India 22 (1): 56 - 59."]}

Despite evidence from anatomy, behavior and genomics indicating that the sense of smell in turtles is important, our understanding of chemical communication in this group is still rudimentary. Our aim was to describe the microanatomy of mental glands (MGs) in a freshwater turtle,Mauremys leprosa(Geoemydidae), and to assess the chemical composition of their secretions with respect to variation among individuals and between sexes. MGs are paired sac-like organs on the gular region of the neck and are dimorphic in this species with males having fully functional holocrine glands while those of females appear non-secretory and vestigial. In adult males, the glandular epithelium of the inner portion of the gland provides exocytotic products as well as cellular debris into the lumen of the gland. The contents of the lumen can be secreted through the narrow duct portion of the gland ending in an orifice on the surface of the skin. Females have invaginated structures similar in general outline to male glands, but lack a glandular epithelium. Using gas chromatography coupled to mass spectrometry, we identified a total of 61 compounds in mental gland secretions, the most numerous being carboxylic acids, carbohydrates, alkanes, steroids and alcohols. The number of compounds per individual varied widely (mean (median) ± SD = 14.54 (13) ± 8.44; min = 3; max = 40), but only cholesterol was found in all samples. We found that the relative abundances of only six chemicals were different between the sexes, although males tended to have larger amounts of particular compounds. Although the lipid fraction of mental gland secretions is rich in chemical compounds, most occur in both sexes suggesting that they are metabolic byproducts with no role in chemical signaling. However, the relative amounts of some compounds tended to be higher in males, with significantly larger amounts of two carboxylic acids and one steroid, suggesting their putative involvement in chemical communication.

Nowadays, the living species of the terrapin genus Mauremys (Testudinoidea: Geoemydidae) are mostly found in eastern Asia, but three of them inhabit the Western Palearctic ecozone. In Italy, occurrences of living individuals of Mauremys are interpreted as records of alien species; however, a growing fossil record demonstrates that this genus has inhabited Italy as recently as the Late Pleistocene. We report on a new fossil specimen of Mauremys from the Upper Pliocene (Piacenzian) marginal-marine deposits of Tuscany (central Italy). This find, consisting of a partial plastron and a loose neural, represents the second authentic report of Mauremys from the Italian Pliocene, as well as the first one from the Piacenzian of Italy. Therefore, it is a significant fossil that fills a gap in the chronostratigraphic distribution of Italian fossil Mauremys, helping – together with the Lower Pliocene holotype of Mauremys portisi – to bridge the rich Miocene and Pleistocene segments of this record. Moreover, two unusual scars observed on the external surface of the studied plastron are here referred to the ichnospecies Thatchtelithichnus holmani. These traces represent one of the few records worldwide of this rarely identified ichnospecies, as well as its geologically youngest published occurrence. Hypotheses regarding the origin of the Thatchtelithichnus traces are reevaluated, and an origin as attachment scars of aquatic ectoparasites (possibly ticks, leeches, or flukes) is reaffirmed as probable in cases of traces occurring on the exterior of the plastral bones of turtles.

Assam Roofed Turtle, Pangshura sylhetensis is an endangered and least studied species endemic to India and Bangladesh. The genomic feature of P. sylhetensis mitogenome is still anonymous to the scientific community. The present study decodes the first complete mitochondrial genome of P. sylhetensis (16,568 bp) by using next-generation sequencing. This de novo assembly encodes 13 Protein-coding genes (PCGs), 22 transfer RNAs (tRNAs), two ribosomal RNAs (rRNAs), and one control region (CR). Most of the genes were encoded on the majority strand, except NADH dehydrogenase subunit 6 (nad6) and eight tRNAs. Most of the PCGs were started with an ATG initiation codon, except for Cytochrome oxidase subunit 1 (cox1) and NADH dehydrogenase subunit 5 (nad5) with GTG. The study also found the typical cloverleaf secondary structure in most of the tRNA genes, except for serine (trnS1) with lack of conventional DHU arm and loop. Both, Bayesian and Maximum-likelihood topologies showed distinct clustering of all the Testudines species with their respective taxonomic ranks and congruent with the previous phylogenetic hypotheses (Pangshura and Batagur sister taxa). Nevertheless, the mitogenomic phylogeny with other amniotes corroborated the sister relationship of Testudines with Archosaurians (Birds and Crocodilians). Additionally, the mitochondrial Gene Order (GO) analysis indicated that, most of the Testudines species showed plesiomorphy with typical vertebrate GO.