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ABSTRACT: The West Asian stripe-necked terrapin Mauremys
caspica is widespread throughout the Middle East—a region
for which only few phylogeographic studies are available.
Due to landscape alteration, pollution and intensification of
water management, M. caspica is increasingly threatened.
However, genetic diversity among and within populations is
poorly known, impeding the identification of management
units. Using a nearly rangewide sampling, we analyzed 14
microsatellite loci and mtDNA sequences in order to gain
insight into the population structure and history of M. caspica.
In agreement with a previous study, we found two clusters of
mitochondrial haplotypes, with one cluster distributed in the
east and the other in the west of the range. However, our
microsatellite data suggested a more pronounced geographical
structuring. When null alleles were coded as recessive
with STRUCTURE 2.3.2, three clusters were revealed, with one cluster matching roughly the range of the western mitochondrial
cluster, and the composite ranges of the two other
microsatellite clusters correspond to the distribution of the
eastern mitochondrial cluster. Naïve STRUCTURE analyses
without correction for null alleles were congruent with respect
to the two eastern microsatellite clusters, but subdivided the
western cluster into two units, with an additional geographical
divide corresponding to the ‘Anatolian diagonal’—a wellknown
high mountain barrier impeding exchange between
western and eastern taxa. In naïve analyses, the westernmost
microsatellite cluster (from Central Anatolia) is quite isolated
from the others, and its distinctness is also supported by
fixation indices resembling the values among the other three
clusters. One of the two eastern clusters is distributed in the
Caucasus region plus Iran, and terrapins from Saudi Arabia
and Bahrain constitute the second eastern cluster, supporting
the view that these endangered populations are native.
Coalescent-based analyses of our microsatellite data reveal
for all four clusters bottlenecks 4,000–20,000 years ago, suggesting
that climatic fluctuations of the Late Pleistocene and
Holocene played an important role in shaping current genetic
diversity. We propose that each of the four identified clusters,
including the Central Anatolian one, should be treated as a
distinct management unit. The presence of non-native terrapins
in the animal trade of Bahrain highlights the danger of
genetic pollution of the endangered Arabian populations.
Further sampling is needed to elucidate the situation in
southern and central Iran and Iraq. Our results confirm that
genetic data do not support the validity of any of the three
morphologically defined subspecies of M. caspica, and we
propose that their usage be abandoned.
Organisms Diversity & Evolution 06/2012; · 1.65 Impact Factor
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Uwe Fritz,
Daniela Guicking,
Hajigholi Kami,
Marine Arakelyan,
Markus Auer,
Dinçer Ayaz,
César Ayres Fernández,
Andrey G Bakiev,
Antonia Celani,
Georg Džuki,
Soumia Fahd,
Peter Havaš,
Ulrich Joger,
Viner F Khabibullin, Lyudmila F Mazanaeva,
Pavel Široký,
Sandro Tripepi,
Aitor Valdeón Vélez,
Guillermo Velo Antón,
Michael Wink
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ABSTRACT: Based on more than 1100 samples of Emys orbicularis and E. trinacris, data on mtDNA diversity and distribution of haplotypes are provided, including for the first time data for Armenia, Georgia, Iran, and the Volga, Ural and Turgay River Basins of Russia and Kazakhstan. Eight mitochondrial lineages comprising 51 individual haplotypes occur in E. orbicularis, a ninth lineage with five haplotypes corresponds to E. trinacris. A high diversity of distinct mtDNA lineages and haplotypes occurs in the south, in the regions where putative glacial refuges were located. More northerly parts of Europe and adjacent Asia, which were recolonized by E. orbicularis in the Holocene, display distinctly less variation; most refuges did not contribute to northern recolonizations. Also in certain southern European lineages a decrease of haplotype diversity is observed with increasing latitude, suggestive of Holocene range expansions on a smaller scale. The mitochondrial cytochrome b gene (cyt b) became a frequently used marker for infer-ring phylogeography in reptiles (e.g. Brown and Pestano, 1998; Burbrink et al., 2000; Carranza et al., 2000, 2002; Surget-Groba et al., 2001; Harris et al., 2002; Austin et al., 2003; Pod-nar et al., 2005; Poulakakis et al., 2005; Fritz et al., 2006a) and several studies on Euro-pean pond turtles (Emys orbicularis, Emys tri-nacris) were based on this marker gene. While Lenk et al. (1999) provided a nearly range-wide phylogeography, their study suffered from a patchy sampling for many parts of the range
Amphibia-Reptilia 01/2007; 28:418-426. · 1.06 Impact Factor
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ABSTRACT: Tortoises of the Testudo graeca complex inhabit a patchy range that covers part of three continents (Africa, Europe, Asia). It extends approximately 6500 km in an east-west direction from eastern Iran to the Moroccan Atlantic coast and about 1600 km in a north-south direction from the Danube Delta to the Libyan Cyrenaica Peninsula. Recent years have seen a rapid increase of recognized taxa. Based on morphological investigations, it was suggested that this group consists of as many as 20 distinct species and is paraphyletic with respect to T. kleinmanni sensu lato and T. marginata. Based on samples from representative localities of the entire range, we sequenced the mitochondrial cytochrome b gene and conducted nuclear genomic fingerprinting with ISSR PCR. The T. graeca complex is monophyletic and sister to a taxon consisting of T. kleinmanni sensu lato and T. marginata. The T. graeca complex comprises six well-supported mtDNA clades (A-F). Highest diversity is found in the Caucasian Region, where four clades occur in close neighbourhood. This suggests, in agreement with the fossil record, the Caucasian Region as a radiation centre. Clade A corresponds to haplotypes from the East Caucasus. It is the sister group of another clade (B) from North Africa and western Mediterranean islands. Clade C includes haplotypes from western Asia Minor, the southeastern Balkans and the western and central Caucasus Region. Its sister group is a fourth, widely distributed clade (D) from southern and eastern Asia Minor and the Levantine Region (Near East). Two further clades are distributed in Iran (E, northwestern and central Iran; F, eastern Iran). Distinctness of these six clades and sister group relationships of (A + B) and (C + D) are well-supported; however, the phylogeny of the resulting four clades (A + B), (C + D), E and F is poorly resolved. While in a previous study (Fritz et al., 2005a) all traditionally recognized Testudo species were highly distinct using mtDNA sequences and ISSR fingerprints, we detected within the T. graeca complex no nuclear genomic differentiation paralleling mtDNA clades. We conclude that all studied populations of the T. graeca complex are conspecific under the Biological Species Concept. There is major incongruence between mtDNA clades and morphologically defined taxa. Morphologically well-defined taxa, like T. g. armeniaca or T. g. floweri, nest within clades comprising also geographically neighbouring, but morphologically distinctive populations of other taxa (clade A: sequences of morphologically similar tortoises of the same subspecies (T. g. ibera sensu stricto or T. g. ibera sensu lato) scatter over two or three genetically distinct clades (A, C or A, C, D, respectively). This implies that pronounced morphological plasticity, resulting in phenotypes shaped by environmental pressure, masks genetic differentiation. To achieve a more realistic taxonomic arrangement reflecting mtDNA clades, we propose reducing the number of T. graeca subspecies considerably and regard in the eastern part of the range five subspecies as valid (T. g. armeniaca, T. g. buxtoni, T. g. ibera, T. g. terrestris, T. g. zarudnyi). As not all North African taxa were included in the present study, we refrain from synonymizing North African taxa with T. g. graeca (mtDNA clade B) that represents a further valid subspecies.
Amphibia-Reptilia 01/2007; 28:97-121. · 1.06 Impact Factor
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ABSTRACT: Tortoises of the Testudo graeca complex inhabit a patchy range that covers part of three continents (Africa, Europe, Asia). It extends approximately 6500 km in an east-west direction from eastern Iran to the Moroccan Atlantic coast and about 1600 km in a north-south direction from the Danube Delta to the Libyan Cyrenaica Peninsula. Recent years have seen a rapid increase of recognized taxa. Based on morphological investigations, it was suggested that this group consists of as many as 20 distinct species and is paraphyletic with respect to T. kleinmanni sensu lato and T. marginata. Based on samples from representative localities of the entire range, we sequenced the mitochondrial cytochrome b gene and conducted nuclear genomic fingerprinting with ISSR PCR. The T. graeca complex is monophyletic and sister to a taxon consisting of T. kleinmanni sensu lato and T. marginata. The T. graeca complex comprises six well-supported mtDNA clades (A-F). Highest diversity is found in the Caucasian Region, where four clades occur in close neighbourhood. This suggests, in agreement with the fossil record, the Caucasian Region as a radiation centre. Clade A corresponds to haplotypes from the East Caucasus. It is the sister group of another clade (B) from North Africa and western Mediterranean islands. Clade C includes haplotypes from western Asia Minor, the southeastern Balkans and the western and central Caucasus Region. Its sister group is a fourth, widely distributed clade (D) from southern and eastern Asia Minor and the Levantine Region (Near East). Two further clades are distributed in Iran (E, northwestern and central Iran; F, eastern Iran). Distinctness of these six clades and sister group relationships of (A + B) and (C + D) are well-supported; however, the phylogeny of the resulting four clades (A + B), (C + D), E and F is poorly resolved. While in a previous study (Fritz et al., 2005a) all traditionally recognized Testudo species were highly distinct using mtDNA sequences and ISSR fingerprints, we detected within the T. graeca complex no nuclear genomic differentiation paralleling mtDNA clades. We conclude that all studied populations of the T. graeca complex are conspecific under the Biological Species Concept. There is major incongruence between mtDNA clades and morphologically defined taxa. Morphologically well-defined taxa, like T. g. armeniaca or T. g. floweri, nest within clades comprising also geographically neighbouring, but morphologically distinctive populations of other taxa (clade A: T. g. armeniaca, T. g. ibera, T. g. pallasi ; clade D: T. g. anamurensis, T. g. antakyensis, T. g. floweri, T. g. ibera, T. g. terrestris), while sequences of morphologically similar tortoises of the same subspecies (T. g. ibera sensu stricto or T. g. ibera sensu lato) scatter over two or three genetically distinct clades (A, C or A, C, D, respectively). This implies that pronounced morphological plasticity, resulting in phenotypes shaped by environmental pressure, masks genetic differentiation. To achieve a more realistic taxonomic arrangement reflecting mtDNA clades, we propose reducing the number of T. graeca subspecies considerably and regard in the eastern part of the range five subspecies as valid (T. g. armeniaca, T. g. buxtoni, T. g. ibera, T. g. terrestris, T. g. zarudnyi). As not all North African taxa were included in the present study, we refrain from synonymizing North African taxa with T. g. graeca (mtDNA clade B) that represents a further valid subspecies.
Amphibia-Reptilia 12/2006; 28(1):97-121. · 1.06 Impact Factor
