We analyse phylogeny, systematics and biogeography of slider turtles (Trachemys spp.) using sequence data of four mitochondrial genes (3242 bp) and five nuclear loci (3396 bp) of most South American and southern Central American taxa and representatives of northern Central American, West Indian and North American slider species (16 species and subspecies) and allied North American species (genera Chrysemys, Deirochelys, Graptemys, Malaclemys, Pseudemys). By applying maximum likelihood, relaxed molecular clock and ancestral range analyses, we provide evidence for two successive colonizations of South America by slider turtles. In addition, we show that the current species delineation of Central and South American slider turtles is incorrect. Our data suggest that Trachemys grayi is a distinct polytypic species that embraces, besides the nominotypical subspecies, T. g. emolli and T. g. panamensis. Trachemys ornata is also polytypic with the subspecies T. o. ornata, T. o. callirostris, T. o. cataspila, T. o. chichiriviche and T. o. venusta. Moreover, T. adiutrix should be regarded as a subspecies of T. dorbigni. All studied Trachemys species are inferred to have originated in the Late Miocene to Early Pliocene. The ancestor of the two subspecies of T. dorbigni colonized South America most probably prior to the establishment of the land bridge connecting Central and South America, whereas the two South American subspecies of T. ornata represent a younger independent immigration wave from Central America.
Hermann's tortoise (Testudo hermanni), the best‐known western Palaearctic tortoise species, has a rare natural distribution pattern comprising the Mediterranean areas of the Iberian, Apennine, and Balkan Peninsulas, as well as Sicily, Corsica and Sardinia. The western part of this range is traditionally considered habitat for T. h. hermanni, while T. h. boettgeri occurs in the Balkans. Taxonomy of this tortoise has been challenged in recent years, with the two subspecies being considered full species and the central Dalmatian populations of T. h. boettgeri being considered a third species, T. hercegovinensis. Using an mtDNA fragment approximately 1150 bp long (cytochrome b gene and adjacent portion of tRNA‐Thr gene), we investigated mtDNA diversity with regard to contrasting concepts of two subspecies or three species. Seven closely related haplotypes were identified from the western Mediterranean and 15 different, in part much‐differentiated, haplotypes from the Balkans. Western Mediterranean haplotypes differ from Balkan haplotypes in 16–42 mutation steps. One to seven mutation steps occur within western Mediterranean populations. Balkan haplotypes, differing in 1−37 nucleotides, group in parsimony network analysis into three major assemblages that display, in part, a similar degree of differentiation to that of western Mediterranean haplotypes relative to Balkan haplotypes. Rates of sequence evolution are different in both regions, and low divergence, palaeogeography and the fossil record suggest a slower molecular clock in the western Mediterranean. While monophyly in western Mediterranean haplotypes is well‐supported, conflicting evidence is obtained for Balkan haplotypes; maximum parsimony supports monophyly of Balkan haplotypes, but other phylogenetic analyses (Bayesian, ML, ME) indicate Balkan haplotypes could be paraphyletic with respect to the western Mediterranean clade. These results imply a process of differentiation not yet complete despite allopatry in the western Mediterranean and the Balkans, and suggest all populations of T. hermanni are conspecific. In the western Mediterranean no clear geographical pattern in haplotype distribution is found. Distribution of Balkan haplotypes is more structured. One group of similar haplotypes occurs in the eastern Balkans (Bulgaria, Republic of Macedonia, Romania and the Greek regions Evvia, Macedonia, Peloponnese, Thessaly and Thrace). Two distinct haplotypes, differing in eight to nine mutation steps from the most common haplotype of the first group, are confined to the western slope of the Taygetos Mts. in the Peloponnese. Yet another group, connected over between four and 23 mutation steps with haplotypes of the eastern Balkan group, occurs along the western slope of the Dinarid and Pindos Mts. (Istria, Dalmatia, western Greece). Taygetos haplotypes are nested within other haplotypes in all phylogenetic analyses and support for monophyly of the other Balkan groups is at best weak. We conclude that using the traditional two subspecies model should be contin...
Phylogeny and taxonomy of endangered South and South-east Asian freshwater turtles elucidated by mtDNA sequence variation (Testudines: Geoemydidae: Batagur, Callagur, Hardella, Kachuga, 36 ,[429][430][431][432][433][434][435][436][437][438][439][440][441][442] Using DNA sequences of the mitochondrial cytochrome b gene, we investigated phylogeny and taxonomy of South and South-east Asian turtles of all species and subspecies of the genera Batagur, Callagur, Hardella, Kachuga and Pangshura. We found three major clades: (i) a moderately to well-supported clade containing all large riverine species assigned so far to Batagur, Callagur and Kachuga ; (ii) a well-supported monophylum comprising the four Pangshura species; and (iii) Hardella that could constitute either the sister-taxon of Pangshura or of a clade comprising Batagur, Callagur, Kachuga and Pangshura. The genus Kachuga is clearly polyphyletic. Therefore, we recommend placing all Batagur, Callagur and Kachuga species in one genus. According to the International Code of Zoological Nomenclature Batagur Gray, 1856, being originally erected at higher rank, takes precedence over the simultaneously published name Kachuga Gray, 1856, and the younger name Callagur Gray, 1870, resulting in an expanded genus Batagur. Indonesian and Malaysian Batagur baska proved to be highly distinct from our sequences of this species from the Sundarbans (Bangladesh, adjacent India), suggesting that a previously unidentified species is involved. This finding is of high conservation relevance in the critically endangered B. baska. The currently recognized subspecies within Hardella thurjii , Pangshura smithii and P. tentoria do not correspond well with mtDNA clades. Considering that the two subspecies of H. thurjii are likely to be based only on individual ontogenetic differences, we propose abandoning the usage of subspecies within H. thurjii . In the Ghaghra River, Uttar Pradesh (India) we detected shared haplotypes in P. smithii and P. tentoria, implying that the unusual morphological characters of the Ghaghra River population of P. tentoria could be the result of interspecific hybridization. Peter Praschag, Am Katzelbach 98, A-8054 Graz, Austria.
rfer, A. K. (2010). Molecular phylogeny of African hinged and helmeted terrapins (Testudines: Pelomedusidae: Pelusios and Pelomedusa). -Zoologica Scripta, 40, 115-125.With 18 currently recognised species, Pelusios is one of the most speciose chelonian genera worldwide, even though the taxonomy of some species is contentious. Recent investigations suggested that the closely related, but morphologically distinct genus Pelomedusa is paraphyletic with respect to Pelusios, and that Pelomedusa consists of nine deeply divergent lineages. Using three mitochondrial and three nuclear DNA fragments (2054 bp mtDNA, 2025 bp nDNA), we examined for the first time the phylogeny of Pelusios by molecular means. Our analyses included all Pelusios species, except the probably extinct P. seychellensis, as well as the nine Pelomedusa lineages. The results showed that Pelusios and Pelomedusa are reciprocally monophyletic. Limited sampling of Pelusios species and homoplasy introduced by remote outgroups most likely explain the paraphyly of Pelomedusa in previous studies. The distinctiveness of most Pelusios species was confirmed, but none of the currently recognised species groups within Pelusios was monophyletic. In Pelusios rhodesianus and P. sinuatus distinct genetic lineages were discovered, suggestive of cryptic taxa. In contrast, the recognition of the weakly differentiated P. castaneus and P. chapini as full species is doubtful, as is the validity of the Malagasy and Seychellois subspecies of P. castanoides. GenBank sequences of P. williamsi were nested within P. castaneus, but the morphological distinctiveness of the two species makes it likely that the GenBank sequences (derived from a turtle from the pet trade) are misidentified. Divergence among the distinct genetic lineages of Pelomedusa equals or exceeds the differences among Pelusios species, supporting the view that Pelomedusa is a species complex.
In the present study, we use mtDNA sequence data (cyt b gene) in combination with nuclear DNA sequences (C‐mos, Rag2 genes, R35 intron), nuclear genomic fingerprints (ISSR) and morphological data to reveal species diversity within the Southeast Asian leaf turtle genus Cyclemys, a morphologically difficult group comprising cryptic species. Two morphologically distinct major groupings exist, a yellow‐bellied species group with three taxa (Cyclemys atripons, C. dentata, C. pulchristriata) and a dark‐bellied species group. The latter contains besides the morphologically variable C. oldhamii three additional new species (C. enigmatica n. sp., C. fusca n. sp., C. gemeli n. sp.). According to mtDNA data, C. fusca and C. gemeli constitute with high support the sister group of a clade comprising all other species, indicating that the dark‐bellied species are not monophyletic, despite morphological similarity. mtDNA sequences of C. enigmatica, being highly distinct in nuclear genomic markers, do not differ from the sympatric C. dentata, suggesting that the original mitochondrial genome of C. enigmatica was lost due to introgressive hybridization. Morphological discrimination of Cyclemys species is possible using multivariate methods. However, gross morphology of most dark‐bellied species on the one hand and of C. atripons and C. pulchristriata on the other is so similar that reliable species determination is only possible when genetic markers are used. The high diversity within Cyclemys requires revision of the IUCN Red List Categories for leaf turtles because the former assessment was based on the wrong assumption that in the entire range of the genus occurs only a single species.
Pelodiscus is one of the most widely distributed genera of softshell turtles, ranging from south-eastern Siberia and Korea over central and southern China to Vietnam. Economically, Pelodiscus are the most important chelonians of the world and have been bred and traded in high numbers for centuries, resulting in many populations established outside their native range. Currently, more than 300 million turtles per year are sold in China alone, and the bulk of this figure comprises farmed Pelodiscus. Due to easy availability, Pelodiscus also constitutes a model organism for physiological and embryological investigations. Yet, diversity and taxonomy of Pelodiscus are poorly understood and a comprehensive investigation using molecular tools has never been published. Traditionally, all populations were assigned to the species P. sinensis (Wiegmann, 1834); in recent years up to three additional species have been recognized by a few authors, while others have continued to accept only P. sinensis. In the present study, we use trade specimens and known-locality samples from Siberia, China, and Vietnam, analyze 2,419 bp of mtDNA and a 565-bp-long fragment of the nuclear C-mos gene to elucidate genetic diversity, and compare our data with sequences available from GenBank. Our findings provide evidence for the existence of at least seven distinct genetic lineages and suggest interbreeding in commercial turtle farms. GenBank sequences assigned to P. axenaria (Zhou, Zhang & Fang, 1991) are highly distinct. The validity of P. maackii (Brandt, 1857) from the northernmost part of the genus' range is confirmed, whereas it is unclear which names should be applied to several taxa occurring in the central and southern parts of the range. The diversity of Pelodiscus calls for caution when such turtles are used as model organisms, because the respective involvement of more than a single taxon could lead to irreproducible and contradictory results. Moreover, our findings reveal the need for a new assessment of the conservation status of Pelodiscus. While currently all taxa are subsumed under 'P. sinensis' and listed as 'vulnerable' by the IUCN Red List of Threatened Species, some could actually be endangered or even critically endangered.
In recent years many cases of hybridization and introgression became known for chelonians, requiring a better understanding of their speciation mechanisms. Phylogeographic investigations offer basic data for this challenge. We use the sister species Mauremys caspica and M. rivulata, the most abundant terrapins in the Near and Middle East and South‐east Europe, as model. Their phylogeographies provide evidence that speciation of chelonians fits the allopatric speciation model, with both species being in the parapatric phase of speciation, and that intrinsic isolation mechanisms are developed during speciation. Hybridization between M. caspica and M. rivulata is very rare, suggesting that the increasing numbers of hybrids in other species are caused by human impact on environment (breakdown of ecological isolation). Genetic differentiation within M. caspica and M. rivulata resembles the paradigm of southern genetic richness and northern purity of European biota. However, in west Asia this pattern is likely to reflect dispersal and vicariance events older than the Holocene. For M. caspica three distinct Pleistocene refuges are postulated (Central Anatolia, south coast of Caspian Sea, Gulf of Persia). Morphologically defined subspecies within M. caspica are not supported by genetic data. This is one of the few studies available about the phylogeography of west and central Asian species.
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