Uwe Fritz and Anja Schulze
Uwe Fritz, Mario Vargas-Ramírez and Pavel Široký
We re-examine the phylogenetic position of Pelusios williamsi by merging new sequences with an earlier published data set of all Pelusios species, except the possibly extinct P. seychellensis, and the nine previously identified lineages of the closely allied genus Pelomedusa (2054 bp mtDNA, 2025 bp nDNA). Furthermore, we include new sequences of Pelusios broadleyi, P. castanoides, P. gabonensis and P. marani. Individual and combined analyses of the mitochondrial and nuclear data sets indicate that P. williamsi is sister to P. castanoides, as predicted by morphology. This provides evidence for the misidentification of GenBank sequences allegedly representing P. williamsi. Such mislabelled GenBank sequences contribute to continued confusion, because only the original submitter can revise their identification; an impractical procedure impeding the rectification of obvious mistakes. We recommend implementing another option for revising taxonomic identifications, paralleling the century-old best practice of natural history museums for new determinations of specimens. Within P. broadleyi, P. gabonensis and P. marani, there is only shallow genetic divergence, while some phylogeographic structuring is present in the wide-ranging species P. castaneus and P. castanoides.
Markus Auer, Uwe Fritz and Maxim Ludwig
Using radiographies of manus and pes of 133 geoemydid terrapins of nine highly aquatic species (genera Batagur, Callagur, Hardella, Kachuga, Orlitia, Pangshura), four aquatic Rhinoclemmys species, three semiterrestrial Heosemys species, and three terrestrial Rhinoclemmys species, we confirm a correlation between mode of life and phalangeal formulae. Terrestrial geoemydid species tend to have lost phalanges or entire digits, while highly aquatic species with extensive toe webbing generally have retained the full phalangeal number in manus and pes. Phalangeal formulae of species that are not so strictly adapted to aquatic environments, and having less extensive toe webbing, are intermediate, like in the studied semiterrestrial species.
Ellen Hitschfeld, Markus Auer and Uwe Fritz
We compared variation in phalangeal and carpal morphology of the Central Asian Testudo horsfieldii, a burrow-digging tortoise species, with its sister taxon T. hermanni, a Mediterranean species without extensive digging behaviour. Thirty-two Testudo horsfieldii kazachstanica shared the same phalangeal formula (0-2-2-1-1). The distal carpal 1 and metacarpal I as well as the pisiform were consistently lacking, sometimes also the medial centrale. Phalangeal morphology was more variable in Testudo hermanni hermanni. In 29 specimens four phalangeal formulae were found that differed with respect to the reduction of digit 1 (1-2-2-2-1, M-2-2-2-1, D-2-2-2-1, 0-2-2-2-1). The pisiform develops late in ontogeny and is lacking in very most juveniles and subadults. In contrast to T. h. kazachstanica, the medial centrale is always present. In both T. h. kazachstanica and T. h. hermanni carpalia increasingly fuse with age and size. The underlying morphological patterns differ however. The extreme character state in aged T. h. kazachstanica is one large solid bone element, formed by the fused intermedium, ulnare and both centralia. Aged T. h. hermanni have, in contrast, two separate larger carpal elements, one formed by the fused lateral and medial centralia and the other by the fused distal carpalia 1 and 2; the intermedium and ulnare never fuse with one another or with other carpalia. While a partial or complete loss of digit 1 seems to be characteristic for all Testudo species, we propose that the extensive fusion of carpal elements in T. horsfieldii is correlated with its natural history because a rigid manus could be advantageous for burrow-digging. Also the reduction of digit 4 (one phalanx present), a rare character among testudinids and not occurring in other Testudo species, could be linked with its mode of life.
Maria Dimaki, Anna Hundsdörfer and Uwe Fritz
Based on mitochondrial 16S rRNA sequences, we suggest that the founder individuals of the introduced Greek population of Chamaeleo africanus originated in the Nile Delta region of Egypt. In Ch. chamaeleon, we discovered in the eastern Mediterranean new 16S rRNA haplotypes, being highly distinct from previously published western Mediterranean haplotypes. Eastern Mediterranean haplotypes were found in samples from northern Syria, Cyprus, Crete, Samos, Malta and Tunisia. The occurrence of an eastern Mediterranean haplotype in Tunisia and of distinct haplotypes in Morocco could argue for a phylogeographic break in northwestern Africa.
Uwe Fritz, Markus Auer and Antje Petzold
Thirty-six complete skeletons, three shells, and x-rays of the extremities of 32 additional turtles of the Cuora galbinifrons complex have been compared with 38 other geoemydid species from 19 genera. Cuora bourreti differs from C. galbinifrons and C. picturata by a lost phalanx in the fourth finger and fourth toe. Individuals with a shell shape intermediate between C. bourreti and C. galbinifrons, as found on Hainan Island (China), have either the reduced phalangeal formula of C. bourreti (manus: 2-3-3-2-2, pes: 2-3-3-2-1) or the complete number of phalanges (manus: 2-3-3-3-2, pes: 2-3-3-3-1). Only in C. flavomarginata did we also register a lost phalanx in the fourth digit of manus and pes; in the pes of C. mouhotii the same character state may occur. In C. flavomarginata the fifth digit of the pes is also lacking. Some other terrestrial and semiterrestrial geoemydids (Cuora mccordi, Heosemys spinosa, and in part C. mouhotii and Leucocephalon yuwonoi) display a similar pattern of phalangeal reduction, resulting in the loss of the fifth digit of the pes. Likewise, in tortoises (Testudinidae), a further group of terrestrial chelonians, and the terrestrial turtle genus Terrapene (Emydidae) the loss of phalanges or complete digits is known to occur. Malayemys subtrijuga, Morenia petersi, Pangshura smithii and Siebenrockiella crassicollis differ from all other studied geoemydid taxa by an additional phalanx in the fifth digit of the manus (2-3-3-3-3); one P. smithii has on one body side three phalanges in the fifth digit of the pes (2-3-3-3-3). These are highly aquatic turtles with extensive toe webbing. Probably, longer digits (and thus a higher phalangeal number) are a favorable prerequisite for swimming while phalangeal loss seems to be the consequence of walking.Cuora bourreti and C. picturata have consistently in the bony carapace a very rare character state regarding the articulation of the rib tips with the peripheral plates. In both species the rib tips are intercalated between two peripheral plates in the bridge region. Intercalated rib tips like those in C. bourreti and C. picturata were found only in C. m. mouhotii, but not in the southern subspecies C. mouhotii obsti. In C. galbinifrons and all other geoemydid taxa studied the rib tips articulate on the bridge within the underlying peripheral and not between two peripherals. In turtles morphologically intermediate between C. bourreti and C. galbinifrons both characters states are found. This suggests that such individuals are hybrids or intergrades. Three known-locality specimens from Hainan Island display both extremes and an intermediate character state. This, together with external morphology and the occurrence of both phalangeal formulae in approximately the same frequency on that island, argues for genetic introgression of C. bourreti on the Hainan population of C. galbinifrons. We conclude that our findings qualify C. galbinifrons and C. bourreti under the Biological Species Concept as conspecific.
Markus Auer, Uwe Fritz and Susann Richter
The hyoid apparati of 25 geoemydid species in 16 genera (including aquatic, semiterrestrial and terrestrial taxa) are compared, and for Cuora galbinifrons bourreti ontogenetic development of the hypoid apparatus is described. Generally, ossification of the hyoid apparatus increases with age. The majority of terrestrial species maintain cartilaginous or partially cartilaginous hyoid bodies and second branchial horns throughout life however. Also in tortoises (Testudinidae) hyoid body and second branchial horns remain cartilaginous, suggesting a positive correlation between mode of life and hyoid morphology (terrestrial species with partly cartilaginous hyoid apparatus vs. aquatic species with fully ossified hyoid apparatus).
Daiber, Karl-Fritz, Herms, Eilert, Meixner, Uwe and Mühling-Schlapkohl, Markus
Claudia Corti, Uwe Fritz, Heiko Stuckas and Melita Vamberger
Using mtDNA sequences and 12 microsatellite loci, we compare populations of Testudo graeca from Sardinia and North Africa. The observed pattern of almost no differentiation combined with reduced variation in the Sardinian population is consistent with introduction in prehistoric or historic times from what is now Tunisia and neighbouring Algeria. Furthermore, in the light of the recently published recommendation to eradicate the non-native T. graeca from Italy, we review recent studies on the archaeological and fossil record, on the phylogeography and population genetics of the three other chelonian species occurring in Sardinia (Emys orbicularis, T. hermanni, T. marginata). We conclude that the extant Sardinian populations of all four species are not native. However, they are and should be safeguarded under EC law (Council Regulation No 338/97 on the Protection of Species of Wild Fauna and Flora; Flora Fauna Habitat Directive: Appendix IV, Art. 12) because they serve as a back-up for the declining mainland populations. Moreover, these populations constitute an important part of the human-shaped natural heritage of the Mediterranean.