Phylogeny and diversification of mountain vipers (Montivipera, Nilson et al. 2001) triggered by multiple Plio-Pleistocene refugia and high-mountain topography in the Near and Middle East

ArticleinMolecular Phylogenetics and Evolution 101 · May 2016with 938 Reads 
How we measure 'reads'
A 'read' is counted each time someone views a publication summary (such as the title, abstract, and list of authors), clicks on a figure, or views or downloads the full-text. Learn more
Cite this publication

Do you want to read the rest of this article?

Request Full-text Paper PDF
Advertisement
  • ... The topography, in combination with the Plio-Pleistocene climatic oscillations that followed the warm Miocene climate, caused repeated retractions of populations surviving the climatic pessima in the refugial areas and subsequent dispersals and expansions to the areas inhabited in present times (Hewitt, 1999). Besides the critical importance of the main European peninsulas and Near Eastern mountain ranges for the evolution of the biotas in the Western Palearctic (Hewitt, 2011), existence of a dynamic array of numerous, smaller glacial refugia has been suggested to exist during the Pliocene and Pleistocene glacials (Kapli et al., 2013;Bellati et al., 2015;Stümpel et al., 2016;Van Riemsdijk et al., 2017). Slower extinction rates and population dispersals from these smaller refugia are reflected in complex phylogeographic patterns and prominent genetic divergences, even within phenotypically very similar taxa (Gvoždík et al., 2010;Mikulíček et al., 2013;Kornilios, 2017). ...
    ... Slower extinction rates and population dispersals from these smaller refugia are reflected in complex phylogeographic patterns and prominent genetic divergences, even within phenotypically very similar taxa (Gvoždík et al., 2010;Mikulíček et al., 2013;Kornilios, 2017). While the existence of refugia in the main European peninsulas, Eastern Mediterranean, and Caucasus is generally recognized as critical for the evolution of recent diversity of reptiles (Tuniyev, 1995;Stewart et al., 2010), the importance of the eastern parts of Western Palearctic, such as Anatolia, Iranian Mountain ranges, and Ponto-Caspian steppes, has only recently become more appreciated (Gvoždík et al., 2012;Jandzik, Avcı andGvoždík, 2013, 2018;Stümpel et al., 2016;Javanbakht et al., 2017). ...
    ... The impact of the topography on the evolutionary history and presence of C. austriaca is particularly illustrative in Anatolia, where the smooth snake occurs in the mountains of the Anatolian Diagonal, which also forms the southern limit of its distribution. The mountain ranges within the diagonal pass across Anatolia in northeast-southwestern orientation, and have played a well-recognized role of being an isolating barrier as well as speciation center (Nilson et al., 1990;Bilgin, 2011;Jandzik, Avcı and Gvoždík, 2013;Kapli et al., 2013;Bellati et al., 2015;Stümpel et al., 2016). Based on the presence of relatively diversified populations in the mountains of the Anatolian Diagonal and north of it, it is possible that it served as a refugial area for the smooth snake populations with retracted ranges during glacials. ...
    Article
    Full-text available
    The smooth snake, Coronella austriaca, is a common snake species widespread in the Western Palearctic region. It does not form conspicuous morphological variants and, although several evolutionary lineages have been distinguished based on the analyses of the mitochondrial DNA sequences, only two subspecies with very limited distribution have been traditionally recognized. Here we present an mtDNA phylogeography of the species using geographically extended sampling while incorporating biogeographically important areas that have not been analyzed before, such as Anatolia, Crimea, and Iran. We find that the smooth snake comprises 14 distinct phylogenetic clades with unclear mutual relationships, characterized by complex genetic structure and relatively deep divergences; some of them presumably of Miocene origin. In general, the biogeographic pattern is similar to other Western Palearctic reptiles and illustrates the importance of the main European peninsulas as well as the Anatolian mountains, Caucasus, and Alborz Mts. in Iran for the evolution of the present-day diversity. Considerable genetic structure present in the smooth snake populations within these large areas indicates the existence of several regional Plio-Pleistocene refugia that served as reservoirs for dispersal and population expansions after the glacial periods. The current taxonomy of C. austriaca does not reflect the rich genetic diversity, deep divergences, and overall evolutionary history revealed in our study and requires a thorough revision. This will only be possible with an even higher-resolution sampling and integrative approach, combining analyses of multiple genetic loci with morphology, and possibly other aspects of the smooth snake biology.
  • ... The Armenian Viper (Montivipera raddei) has a fragmented distribution that includes eastern Turkey, Armenia ( Fig.1), Azerbaijan, and northwestern Iran (Nilson and Andrén 1986). Gene flow and range expansion are restricted due to their allopatric populations in isolated mountain habitat above 1300 m and unsuitable habitat in the intervening areas (Nilson and Andrén 1986;Stümpel et al. 2016 2015]). Over the past 40 y there has been a steady decline in population numbers: 20-50 specimens/ha in the 1960s to current estimates of 4-10 specimens/ha (Darevsky 1966;Mallow et al. 2003;IUCN. ...
    ... To date, genetic studies have focused on the phylogenetic relationships of the nine species comprising Montivipera and the taxonomic position of the genus within Viperidae (Nilson et al. 1999;Lenk et al. 2001;Stümpel et al. 2016). No studies have examined the population structure of the Armenian Viper or any of the other Montivipera species. ...
    ... Their current distribution has been restricted further by human-mediated habitat alterations. Due to the long-term isolation of the allopatric populations of M. raddei (Yousefi et al. 2015;Stümpel et al. 2016) we also predicted that there would be strong genetic differentiation between the geographically separated Abovian and Shikahogh populations. ...
    Article
    Full-text available
    Armenian Vipers, Montivipera raddei, have a fragmented distribution in portions of eastern Turkey, Azerbaijan, Armenia, and northwestern Iran. They are restricted to mountain habitat above 1300 m and have allopatric populations resulting from climatic oscillations of the Plio-Pleistocene periods. Anthropogenic landscape changes and over-collection for the pet trade have resulted in dramatic population declines over the past four decades. While we have some insights into spatial ecology and habitat use of this species, we know nothing about their population structure. We examined the genetic diversity and population structure of Armenian Vipers inhabiting an agricultural landscape and a recovered-natural landscape separated by 397 km. We used seven microsatellite loci to genotype 63 individuals representing two sampling locations in each of the two sites. There were no indications of population bottlenecks within any of the sampling locations. While we found evidence of inbreeding at one of the locations in the agricultural landscape, the FST value indicates that individuals can still make contact with one another. We found no significant differentiation between sampling locations at the local scale (FST values of 0.03 to 0.006), but highly significant differentiation between the geographically separated populations (FST ranged from 0.14 to 0.20). The Bayesian clustering algorithm STRUCTURE also identified two distinct population clusters, one consisting of the two agricultural sites and the other the two recovered-natural sites. Conservation efforts should focus on maintaining high quality habitat corridors that allow for gene flow and the management of the geographically separated populations as independent genetic units.
  • ... A few taxa have been described in the last 150 y (Ottoman Viper, M. xanthina, Radde's Viper, M. raddei, Lebanon Viper, M. bornmuelleri, and Latifi's Viper, M. latifii), whereas more recent taxonomic research resulted in the description of a number of new taxa (Wagner's Viper, M. wagneri, Bolkar Viper, M. bulgardaghica, Albizona Viper, M. albizona; and Kuhrang Viper, M. kuhrangica). Species delimitation among these taxa has remained controversial (e.g., Andren 1986, 1992;Schätti et al. 1991;Sindaco et al. 2013), but some recent studies provided more clarity on relationships among mountain vipers (Stümpel and Joger 2009;Stümpel 2012;Stümpel et al. 2016). ...
    ... We engaged in a holistic approach and compiled data from all available sources, including annual field excursions and sampling of representative genetic tissues for species delineation, searching all literature references, contacting authors of online photos and reports, questioning locals as well as herpetologists that have been active in Turkey. We here largely follow the taxonomic concept of Stümpel et al. (2016) but expect changes in the future (Freitas et al. 2020). ...
    ... Recently, M. albizona was suggested as a subspecies of the Bolkar Viper (M. bulgardaghica) due to molecular evidence (Stümpel and Joger 2009;Stümpel et al. 2016). Because the distinction between the two subspecies has become blurrier with new, often photographic, material, we apply the subspecies epithet of such specimens listed herein based on its proximity to the historically known distribution, and/or topographically linked habitats. ...
    Article
    Full-text available
    Turkey harbors a high diversity of viperid snakes, many with a high threat level on the International Union for Conservation of Nature (IUCN) Red List, yet perception about even basic topics, such as distributions and conservation statuses, remain poor. We initiated a multi-year project 7 y ago to compensate these shortcomings and present herein dramatically improved information on the status of mountain vipers of central-eastern Anatolia (Asian Turkey): Bolkar Viper (Montivipera b. bulgardaghica), Albizona Viper (M. b. albizona), Wagner's Viper (M. wagneri), and partly Ottoman Viper (M. xanthina). The data originate from our fieldwork and a comprehensive search of all records available, including information from literature, online resources, locals, and herpetological experts. This resulted in 51 new localities, complemented by 36 published records, which were refined with new information, including four corrected/removed records and two records that were combined with new records due to their proximity. We summarized all records with precise information in a supplemented list of 85 localities, which is compared to current literature and the range maps available on the IUCN Red List of Threatened Species, the global standard reference for consultation on range maps and conservation status of species. Consequently, we report on large range extension of > 100 km in all four mountain viper taxa, increase the extent of occurrence for each viper taxon 4-8 times, reduce the distribution gaps between all pairs of parapatric, related, and ecologically similar mountain vipers, and discuss taxa delimitation, putative contact zones and conservation aspects.
  • ... A few taxa have been described in the last 150 y (Ottoman Viper, M. xanthina, Radde's Viper, M. raddei, Lebanon Viper, M. bornmuelleri, and Latifi's Viper, M. latifii), whereas more recent taxonomic research resulted in the description of a number of new taxa (Wagner's Viper, M. wagneri, Bolkar Viper, M. bulgardaghica, Albizona Viper, M. albizona; and Kuhrang Viper, M. kuhrangica). Species delimitation among these taxa has remained controversial (e.g., Andren 1986, 1992;Schätti et al. 1991;Sindaco et al. 2013), but some recent studies provided more clarity on relationships among mountain vipers (Stümpel and Joger 2009;Stümpel 2012;Stümpel et al. 2016). ...
    ... We engaged in a holistic approach and compiled data from all available sources, including annual field excursions and sampling of representative genetic tissues for species delineation, searching all literature references, contacting authors of online photos and reports, questioning locals as well as herpetologists that have been active in Turkey. We here largely follow the taxonomic concept of Stümpel et al. (2016) but expect changes in the future (Freitas et al. 2020). ...
    ... Recently, M. albizona was suggested as a subspecies of the Bolkar Viper (M. bulgardaghica) due to molecular evidence (Stümpel and Joger 2009;Stümpel et al. 2016). Because the distinction between the two subspecies has become blurrier with new, often photographic, material, we apply the subspecies epithet of such specimens listed herein based on its proximity to the historically known distribution, and/or topographically linked habitats. ...
    Article
    Full-text available
    Turkey harbors a high diversity of viperid snakes, many with a high threat level on the International Union for Conservation of Nature (IUCN) Red List, yet perception about even basic topics, such as distributions and conservation statuses, remain poor. We initiated a multi-year project 7 y ago to compensate these shortcomings and present herein dramatically improved information on the status of mountain vipers of central-eastern Anatolia (Asian Turkey): Bolkar Viper (Montivipera b. bulgardaghica), Albizona Viper (M. b. albizona), Wagner’s Viper (M. wagneri), and partly Ottoman Viper (M. xanthina). The data originate from our fieldwork and a comprehensive search of all records available, including information from literature, online resources, locals, and herpetological experts. This resulted in 51 new localities, complemented by 36 published records, which were refined with new information, including four corrected/removed records and two records that were combined with new records due to their proximity. We summarized all records with precise information in a supplemented list of 85 localities, which is compared to current literature and the range maps available on the IUCN Red List of Threatened Species, the global standard reference for consultation on range maps and conservation status of species. Consequently, we report on large range extension of > 100 km in all four mountain viper taxa, increase the extent of occurrence for each viper taxon 4–8 times, reduce the distribution gaps between all pairs of parapatric, related, and ecologically similar mountain vipers, and discuss taxa delimitation, putative contact zones and conservation aspects.
  • ... x. dianae. Molecular phylogeny and variation of genus Montivipera is described by detailed study of Stümpel et al. (2016) and four lineages; Aegean and Greece lineage and Lycia and Taurus were determined in "xanthina" group. Even though they are geographically close, it is stated that specimens from Greece and Thracian part of Turkey do not share the same clade. ...
    ... Even though they are geographically close, it is stated that specimens from Greece and Thracian part of Turkey do not share the same clade. In parallel with Stümpel et al. (2016), Cattaneo (2017) determined morphological differences in Greek Thrace population and described the subspecies M. x. occidentalis. In the Stümpel et al. (2016), all known taxa (except Chios and Leros island populations) were included in the analysis, and the genetic data stated that M. xanthina generate a cryptic species complex with three or four new taxa. ...
    ... In parallel with Stümpel et al. (2016), Cattaneo (2017) determined morphological differences in Greek Thrace population and described the subspecies M. x. occidentalis. In the Stümpel et al. (2016), all known taxa (except Chios and Leros island populations) were included in the analysis, and the genetic data stated that M. xanthina generate a cryptic species complex with three or four new taxa. However, it is also stated that phenotypical distinction (taxonomical distinction) was not possible due to lack of material and morphological characteristics might be helpful for determining these taxa. ...
    Article
    Full-text available
    A new Ottoman viper subspecies, M. xanthina varoli n. subsp., is described from the higher altitudes of Gündoğmuş (Antalya). The new subspecies differs from the other M. xanthina populations by pholidosis; higher number of intercanthals and lower number of subcaudalia. Also, the whiteness between windings or spots on dorsum pattern were observed in new subspecies, similar to the southern populations. Furthermore, the spots on the ventrals became denser in the mid-body and forms darker colorization at the end of body of the males and the tail tips are yellowish-orange or light orange on both sexes.
  • ... In addition, severe ecological conditions and high spatial divergence in most parts of these Mediterranean ecosystems have resulted in the emergence of a substantial degree of recent speciation events 38 , i.e. neo-endemism 43,44 . As a consequence, the Near and Middle East embrace the richest herpetofauna and harbor the highest diversity of true vipers within the Palearctic bioregion 44,45 . ...
    ... Strong isolations through repeated retractions to isolated mountain tops during late Tertiary climatic fluctuations have caused vicariant allopatric speciation and a rapid rate of diversifications in this genus 7,45 . Generally (and from a phylogenetic viewpoint), Montivipera species are categorized into two complexes; Xanthina and Raddei 37 , yet their taxonomy remains debated 45 . The Xanthina complex includes the species M. xanthina, M. wagneri, and M. bornmuelleri with two species-subspecies M. b. albizona and M. b. bulgardaghica. ...
    ... Another key assumption of ENMs is that a species is in equilibrium with the climate 67,73 , meaning that the observed distribution represents the total of suitable habitats of the species. Regarding the evolutionary history of Montivipera, high degrees of new-endemism, isolated distribution to mountain and limited dispersal abilities 45,74 might have eradicated the species from some areas that once were suitable. This could mean that Montivipera species might be isolated from otherwise suitable habitats, thus violating to some degree the equilibrium assumption. ...
    Article
    Full-text available
    Climate change is among the most important drivers of biodiversity decline through shift or shrinkage in suitable habitat of species. Mountain vipers of the genus Montivipera are under extreme risk from climate changes given their evolutionary history and geographic distribution. In this study, we divided all Montivipera species into three phylogenetic-geographic Montivipera clades (PGMC; Bornmuelleri, Raddei and Xanthina) and applied an ensemble ecological niche modelling (ENM) approach under different climatic scenarios to assess changes in projected suitable habitats of these species. Based on the predicted range losses, we assessed the projected extinction risk of the species relative to IUCN Red List Criteria. Our result revealed a strong decline in suitable habitats for all PGMCs (63.8%, 79.3% and 96.8% for Xanthina, Raddei and Bornmuelleri, respectively, by 2070 and under 8.5 RCP scenario) with patterns of altitudinal range shifts in response to projected climate change. We found that the mountains close to the Mediterranean Sea are exposed to the highest threats in the future (84.6 ± 9.1 percent range loss). We also revealed that disjunct populations of Montivipera will be additionally highly isolated and fragmented in the future. We argue that leveraging climate niche projections into the risk assessment provides the opportunity to implement IUCN criteria and better assess forthcoming extinction risks of species.
  • ... Among viperine snakes, the mountain vipers (genus Montivipera) have been the subject of controversial taxonomic revisions due to the great variability and homoplasy of morphological characters. The genus Montivipera includes 2 sister evolutionary lineages morphologically and genetically well differentiated (Stümpel and Joger 2009;Stümpel et al. 2016): the "xanthina-complex" that inhabits Asia Minor, Syria, and Lebanon and the "raddei-complex" which is distributed across north, northwestern, and western Iran, southern half of Armenia, south of Azerbaijan, Nakhchivan, and eastern Turkey (Nilson and Andrén 1986). The Iranian high plateau contains all the 5 known taxa of the raddei-complex ( Figure 1): M. raddei raddei (Boettger 1890), Montivipera raddei kurdistanica (Nilson and Andrén 1986), Montivipera albicornuta (Nilson and Andrén 1986), Montivipera latifii (Mertens et al. 1967), and Montivipera kuhrangica (Rajabizadeh et al. 2011). ...
    ... The Iranian high plateau contains all the 5 known taxa of the raddei-complex ( Figure 1): M. raddei raddei (Boettger 1890), Montivipera raddei kurdistanica (Nilson and Andrén 1986), Montivipera albicornuta (Nilson and Andrén 1986), Montivipera latifii (Mertens et al. 1967), and Montivipera kuhrangica (Rajabizadeh et al. 2011). However, based on mitochondrial and nuclear genes, Stümpel et al. (2016) showed that M. r. kurdistanica and M. albicornuta do not represent monophyletic assemblages that they should be collapsed into M. raddei. ...
    ... The last glacial maximum [LGM, about 21 kyr (thousand years) ago] corresponded, in the Sahara and Middle East, to a period of higher aridity than presently (Blanchet et al. 1997) and conditions prevailing in this region during the Holocene were much more humid than nowadays. Stümpel et al. (2016) revealed the importance of Pleistocene climatic fluctuations and Middle East relief on mountain viper diversification. Thus, the M. raddei-complex constitutes a case study for studying the consequences of Quaternary glacial cycles in terms of conservation for a highly fragmented taxon. ...
  • ... Differentiation between the latter two clades has been initiated probably by a vicariant speciation event (i.e., isolation of Anatolia during the middle and late Miocene; Stümpel et al., 2016). Nevertheless, published works on the fossil viperine snakes from Europe rarely follow these taxonomical changes Rage, 1999, 2002;Rage, 2013;Georgalis et al., 2016;Čerňanský et al., 2017), mainly because the above established recent taxa are based on external morphology and/or mitochondrial and nuclear DNA differences, and obvious diagnostic features of the skeletal morphology are generally missing. ...
    ... The latter is the most widely distributed viperine genus (i.e., from south-eastern Asia, Middle East, south-eastern Europe and north-western Africa; Uetz et al., 2017). Macrovipera and Montivipera, the remaining two genera partitioned from the genus Vipera (s.l.), according to Page 13 of 26 A c c e p t e d M a n u s c r i p t Stümpel et al. (2016), may be of a more recent origin that resulted from allopatric speciation driven by geographic vicariance. Based on the calculations of Stümpel (2012), Macrovipera and Montivipera shared their last common ancestor during the Langhian (15.3 Ma; Stümpel et al., 2016: fig. ...
    ... Macrovipera and Montivipera, the remaining two genera partitioned from the genus Vipera (s.l.), according to Page 13 of 26 A c c e p t e d M a n u s c r i p t Stümpel et al. (2016), may be of a more recent origin that resulted from allopatric speciation driven by geographic vicariance. Based on the calculations of Stümpel (2012), Macrovipera and Montivipera shared their last common ancestor during the Langhian (15.3 Ma; Stümpel et al., 2016: fig. 3), a divergence time that correlates with the long standing isolation of the Anatolian subcontinent according to the lithological-palaeogeographic maps of Rögl (1999) and Popov et al. (2004). ...
    Article
    We report herein the presence of a large viperine snake assigned to Macrovipera sp. from the early Vallesian (MN 9) locality of Cretesti-1 (Moldova, Romania), representing the first occurrence of this genus in the late Miocene of eastern Romania. We use morphological, morphometrical, phylogeographical and palaeobiogeographical data to differentiate Macrovipera from its sister taxon Montivipera and to reconstruct its supposed late Miocene westward migration alongside the northern margins of Eastern and Central Paratethys from its alleged centre of origin in the Middle East. We assess also the presence of this taxon in several middle and late Miocene localities of Romania, Hungary and Austria, an information completely ignored in recent reviews.
  • ... nov. Given the commonly observed biogeographic pattern in this region (Jandzik, Avcı & Gvoždík, 2013;Jandzik et al., 2018;Stümpel et al., 2016;Tamar et al., 2016;Kornilios, 2017), it is also possible that these, likely, isolated populations have diverged from one of these species (or their common ancestor) and represent a yet undiscovered taxon (Zinner, 1972). ...
    ... Although our estimation is only approximate, it coincides with the splits among other snake and lizard taxa from the same region, most notably the montane vipers of Montivipera xanthina complex-M. xanthina clade from western Anatolia and M. bornmuelleri clade from the east and south of the Anatolian Diagonal separated around five to six Mya (Stümpel et al., 2016). However, these authors propose that the valley of Göksu River south-west of the Anatolian Diagonal formed the natural barrier for the vipers adapted to the life in high altitudes, which was probably not true for the eurytopic rat snakes. ...
    Article
    Full-text available
    Background: The rat snake genus Elaphe once comprised several dozens of species distributed in temperate through tropical zones of the New and Old World. Based on molecular-genetic analyses in early 2000s, the genus was split into several separate genera, leaving only 15 Palearctic and Oriental species as its members. One of the three species also occurring in Europe is Elaphe sauromates, a robust snake from the Balkans, Anatolia, Caucasus, Ponto-Caspian steppes, and Levant that has been suspected to be composed of two or more genetically diverse populations. Here, we studied the genetic structure and morphological variation of E. sauromates, aiming to better understand its inter-population relationships and biogeography, and subsequently revise its taxonomy. Methods: We reconstructed the phylogeography and analyzed the genetic structure of E. sauromates populations originating from most of its geographic range using both mitochondrial (COI, ND4) and nuclear (C-MOS, MC1R, PRLR, RAG1) DNA gene fragments. We employed Maximum likelihood and Bayesian inference methods for the phylogenetic tree reconstructions, supplemented with species delimitation methods, analysis of haplotype networks, and calculation of uncorrected p-distances. Morphological variation in 15 metric and 18 meristic characters was studied using parametric univariate tests as well as multivariate general linearized models. In total, we analyzed sequences originating from 63 specimens and morphological data from 95 specimens of E. sauromates sensu lato. Results: The molecular phylogeny identified two clearly divergent sister lineages within E. sauromates, with both forming a lineage sister to E. quatuorlineata. The genetic distance between them (5.80–8.24% in mtDNA) is similar to the distances among several other species of the genus Elaphe. Both lineages are also moderately morphologically differentiated and, while none of the characters are exclusively diagnostic, their combination can be used for confident lineage identification. Here, following the criteria of genetic and evolutionary species concepts, we describe the lineage from eastern Anatolia and parts of the Lesser and Great Caucasus as a new species E. urartica sp. nov. Discussion: Elaphe urartica sp. nov. represents a cryptic species whose ancestors presumably diverged from their common ancestor with E. sauromates around the Miocene-Pliocene boundary. The intraspecific genetic structure indicates that the recent diversity of both species has been predominantly shaped by Pleistocene climatic oscillations, with glacial refugia mainly located in the Balkans, Crimea, and/or Anatolia in E. sauromates and Anatolia and/or the Caucasus in E. urartica sp. nov.
  • ... However, the knowledge gained by proximate cause questions can be incorporated into a higher-level ultimate cause framework by formulating an appropriate evolutionary hypothesis. In this regard, the timecalibrated framework of snake phylogeny and snake venom evolution is well established [12], and the emphasis on a combination of multiple omic approaches and other disciplines of ancestral state reconstruction, such as paleontology, evolutionary ecology, and biogeography [13,14], adds new dimensions to integrative venomics in its ability to put forth compelling, testable, and falsifiable hypotheses about venom biology and evolution across the full taxonomical range of crown caenophidian snakes. ...
  • ... ''refugia-within-refugia") were present within the current range of O. vittatus. Multiple refugia within the current range of O. vittatus have been found for reptiles (Kornilios et al., 2011;Stümpel et al., 2016), insects (Simonato et al., 2007) and plants (Médail and Diadema, 2009). ...
    Article
    Full-text available
    The banded newt (genus Ommatotriton) is widely distributed in the Near East (Anatolia, Caucasus and the Levant) - an understudied region from the perspective of phylogeography. The genus is polytypic, but the number of species included and the phylogenetic relationships between them are not settled. We sequenced two mitochondrial and two nuclear DNA markers throughout the range of Ommatotriton. For mtDNA we constructed phylogenetic trees, estimated divergence times using fossil calibration, and investigated changes in effective population size with Bayesian skyline plots and mismatch analyses. For nuDNA we constructed phylogenetic trees and haplotype networks. Species trees were constructed for all markers and nuDNA only. Species distribution models were projected on current and Last Glacial Maximum climate layers. We confirm the presence of three Ommatotriton species: O. nesterovi, O. ophryticus and O. vittatus. These species are genetically distinct and their most recent common ancestor was dated at ∼25 Ma (Oligocene). No evidence of recent gene flow between species was found. The species show deep intraspecific genetic divergence, represented by geographically structured clades, with crown nodes of species dated ∼8-13 Ma (Miocene to Early Quaternary); evidence of long-term in situ evolution and survival in multiple glacial refugia. While a species tree based on nuDNA suggested a sister species relationship between O. vittatus and O. ophryticus, when mtDNA was included, phylogenetic relationships were unresolved, and we refrain from accepting a particular phylogenetic hypothesis at this stage. While species distribution models suggest reduced and fragmented ranges during the Last Glacial Maximum, we found no evidence for strong population bottlenecks. We discuss our results in in the light of other phylogeographic studies from the Near East. Our study underlines the important role of the Near East in generating and sustaining biodiversity.
  • ... In total 14 taxa of vipers (Viperinae) have been recorded in Turkey belonging to the genera Macrovipera, Montivipera and Vipera. Mountain Vipers (members of the genus Montivipera) consist of a taxonomic species complex and all species in this group are adapted to rocky habitats above 1500 m, except for Montivipera xanthina which is also found at sea level (Stümpel et al., 2016). Two species groups are described in this complex based on morphological characters: the Xanthina species-group and the Raddei species-group (Mallow et al., 2003;Nilson and Andr en, 1986). ...
    Article
    Here we report the first characterization of the endemic Mount Bulgar Viper (Montivipera bulgardaghica) and Radde's mountain viper (Montivipera raddei) venom by a combined approach using intact mass profiling and bottom-up proteomics. The cytotoxicity screening of crude venom as well as isolated serine proteases revealed a high activity against A549 human lung carcinoma cells. By means of intact mass profiling of native and reduced venom we observed basic and acidic phospholipases type A2. Moreover, the analysis revealed snake venom metalloproteases, cysteine-rich secretory proteins, disintegrins, snake venom serine proteases, C-type lectins, a vascular endothelial growth factor and an L-amino acid oxidase.
  • ... Th e north/east-south/west orientation of the diagonal provides the connection between the east-west directed mountains of the Mediterranean and the Caspian Sea ( Médail and Diadema, 2009 ;Ansell et al., 2011 ). Th e Anatolian Diagonal has been shown to serve as a barrier for some animal species ( Gül, 2013 ;Vamberger et al., 2013 ) or as a diversifi cation spot for other groups ( Stümpel et al., 2016 ). However, as Davis (1971) already suggested, the Anatolian Diagonal might have served as a land bridge between east and west for plants. ...
    Article
    Full-text available
    Premise of the study: The Irano-Turanian region harbors three biodiversity hotspots and ∼25% of Brassicaceae species are endemic to the region. Aethionema (∼61 species) is the sister lineage to the core Brassicaceae and occurs mainly in the Irano-Turanian region. The evolutionary important position of Aethionema makes it an ideal reference for broader comparative genetics and genomics. To understand the evolution of Aethionema, and for a broader understanding of crucifer evolution, a time-calibrated phylogenetic tree and biogeographical history of the genus is needed. Methods: Seventy-six plastome coding regions and nuclear rDNA genes, mainly from herbarium material, covering 75% of all Aethionema species, were used to resolve a time-calibrated Aethionema phylogeny. The different clades were characterized based on four morphological characters. The ancestral area of Aethionema was estimated with historical biogeographical analyses. Key results: Three well-supported major clades within Aethionema were resolved. The ancestral area reconstruction and divergence-time estimates are consistent with major dispersal events during the Pliocene from the Anatolian Diagonal. Conclusions: We find that most Aethionema lineages originated along the Anatolian Diagonal, a floristic bridge connecting the east to the west, during the Pliocene. The dispersal of Aethionema correlates with the local geological events, such as the uplift of the Anatolian and Iranian plateaus and the formation of the major mountain ranges of the Irano-Turanian region. Knowing the paleo-ecological context for the evolution of Aethionema, in addition to the other lineages of Brassicaceae, facilitates our broader understanding for trait evolution and species diversification across the Brassicaceae.
  • ... For example, the river Göksu Nehri breaks the Taurus Mountains with a valley bottom < 250 m asl. between the towns Göksu and Silifke in Mersin Province. This river acts as a barrier for the dispersal of montane organisms, e.g., dividing Montivipera populations into an eastern (bulgardaghica) and a western (xanthina)-clade (Stümpel et al., 2016). Further upstream near Hadim, Konya Province, 40 km northeast of the V. a. senliki ssp. ...
    Article
    Full-text available
    We report on a new population of Vipera anatolica from the Geyik Mountain Range in eastern Antalya Province, Turkey. It represents only the second known location, and is situated in a valley about 200 km east from the terra typica at Kohu Dağ in western Antalya Province. We compare both populations and, based on marked differences in morphology, habitat, genetics, and its geographically isolated location, we describe the recently discovered population as a new subspecies. Aspects of ecology, threats, and conservation needs are discussed.
  • ... Recentemente STÜMPEL et al. (2016) hanno assegnato a un "clade greco" le popolazioni di Montivipera xanthina che abitano i territori a nord-est di Alessandropoli (Evros sud-orientale), ma non fanno alcun riferimento alle popolazioni della restante parte della Tracia greca centro-orientale. Per quanto riguarda l'Evros sud-occidentale, cioè l'area considerata in questa nota, in base a quanto detto, anche se presente, Montivipera xanthina sarà rappresentata da pochi esemplari molto localizzati e sparsi (esemplari relitti di un'epoca caratterizzata da una minore antropizzazione), per cui il flusso genico che ne può derivare verso le limitrofe popolazioni dei Rodopi risulterebbe essere molto limitato. ...
    Article
    Full-text available
    SOMMARIO La distribuzione europea di Montivipera xanthina comprende la fascia costiera della Tracia greca centro-orientale (Rodopi sud-orientali ed Evros meridionale). Mentre nell'Evros sud-orientale, cioè nel tratto più a est di questa distribuzione, M. xanthina sembra rappresentare una linea evolutiva ben definita, nella parte centrale di questo vasto territorio, cioè nell'Evros sud-occidentale, questa vipe-ra sembra essere molto rara o assente; ciò comporta che le popolazioni più occidentali (quelle dei Rodopi) si vengono a trovare in condizioni di isolamento o semisolamento, tali da poter indurre la formazione di una nuova sottospecie: Montivipera xanthina occidentalis ssp. nov. Il nuovo taxon si distingue dalle altre sottospecie note per avere la tendenza alla riduzione delle dimensioni e per avere corrispondentemente un più basso numero di intercantali e di file di squame dorsali, sia a metà tron-co (per lo più 21-22) sia nel tratto posteriore del dorso (spesso 15-16). Viene riportato inoltre un elenco delle specie erpetologiche riscontrate nell'area di studio (Evros sud-occidentale, Grecia NE).
  • ... Scinax species, [142]; Hypsiboas species, [143]); reptiles (e.g. Eulamprus species, [144]; Drysdalia species, [145]; Montivipera species, [146]) and mammals (e.g. Myosorex species, [147]; Nannomys, Aethomys, Otomys, Myotomys, Rhabdomys, Mastomys, Saccostomus, Cryptomys and Xerus species, [148]; Mustela species, [149]). ...
    Article
    Full-text available
    The Anseriformes is a well-known and widely distributed bird order, with more than 150 species in the world. This paper aims to revise the classification, determine the phylogenetic relationships and diversification patterns in Anseriformes by exploring the Cyt b, ND2, COI genes and the complete mitochondrial genomes (mito-genomes). Molecular phylogeny and genetic distance analyses suggest that the Dendrocygna species should be considered as an independent family, Dendrocygnidae, rather than a member of Anatidae. Molecular timescale analyses suggests that the ancestral diversification occurred during the Early Eocene Climatic Optimum (58 ~ 50 Ma). Furthermore, diversification analyses showed that, after a long period of constant diversification, the median initial speciation rate was accelerated three times, and finally increased to approximately 0.3 sp/My. In the present study, both molecular phylogeny and diversification analyses results support that Anseriformes birds underwent rapid and recent diversification in their evolutionary history, especially in modern ducks, which show extreme diversification during the Plio-Pleistocene (~ 5.3 Ma). Therefore, our study support that the Plio-Pleistocene climate fluctuations are likely to have played a significant role in promoting the recent diversification for Anseriformes.
  • ... Latifi (Mertens, Darewsky & Klemmer, 1967) and M. albicornuta (Nilsen & Andren, 1985) all are belonging to M. raddei (Boettger, 1890). Another phylogenetic study conducted by Stümpel et al. (2016) suppose that M. kuhrangica and M. latifi are as valid taxa. (Mertens, Darewsky & Klemmer, 1967) Vipera latifii Mertens, Darevsky & Klemmer, 1967: 161-168. ...
    Article
    Pseudocerastes urarachnoides is a fascinating viper and as yet has been reported only in western Iran. An elaborated arachnid-like caudal structure is a unique feature of this viper, hence gives it the common name “Iranian spidertailed viper”. During tail wagging, the structure is reminiscent of a moving spider. Tail movements are used for two different purposes in snakes: defense via tail vibration and hunting via both caudal luring and caudal distraction. Caudal luring in snakes is the wriggling or wagging of the posterior part of tail, in the presence of a potential prey, with conspicuous color pattern while the rest of body is cryptically colored. Previous studies have speculated on the role of caudal structure of P. urarachnoides in hunting. Our 2.5-year study has revealed that development of the structure of the caudal lure is commenced after birth and is linearly correlated to snout-vent length. The caudal lure attracts some species of birds. Caudal luring behavior is carried out both in the presence and absence of birds. The findings are reported for the first time and confirmed by direct observation of undisturbed individuals in the field.
  • ... Viperinae (true vipers) comprises three major clades, with one comprising species predominantly occurring in Africa (the 'African clade'), another including two genera from the Middle-East, but also extending to some African regions ('carpet' and the 'horned' and 'sand' vipers) and a third with lineages radiating in Europe, Middle-East, Africa and Asia ('Eurasian' vipers). The Near and Middle-East are notable for harbouring the highest diversity of viperines within Eurasia (Stümpel et al., 2016). ...
    Chapter
    Vipers are venomous snakes characterised by having two enlarged highly mobile fangs. They comprise more than 300 species belonging to the Viperidae, which in turn includes viperines (or ‘true vipers’), azemiopines and crotalines (or ‘pitvipers’). They first radiated in the Old World more than 50 million years ago and later rapidly diversified in the New World. Several interesting traits emerged during their evolutionary history, including thermosensitive loreal pits and parental care in crotalines and specialised sound production in rattlesnakes. Vipers are currently found in almost every available habitat on earth and occupy diverse ecological niches. Ancestrally, they evolved the ability to eat unusually heavy, bulky prey and exhibit an impressive range of diet specialisations. Although vipers represent an interesting and successful radiation, many species are threatened with extinction and several aspects of their evolutionary history still need investigation.
  • ... nov., and 125 of Z. situla, Fig. 1 and Supplementary Fig. 3); yet, a northern boundary farther inland can't be specified due to the lack of data. At its eastern distribution the Göksu River and the Mut basin may act as phylogeographic barrier, as it is suggested from other montane biota, such as between the mountain vipers Montivipera xanthina and M. bulgardaghica (Stümpel, Rajabizadeh, Avcı, Wüster, & Joger, 2016), or possibly as an eastern range limit for Vipera anatolica (Göçmen, Mebert, Karış, Oğuz, & Ursenbacher, 2017). ...
    Article
    Based on morphological characteristics, two subspecies of the Transcaucasian rat snake (Zamenis hohenackeri) are currently recognized, namely Z. h. tauricus and Z. h. hohenackeri. Both subspecies are repeatedly considered to be conspecific colour morphs, or have even been confused with Z. situla. Although, few studies involved the Transcaucasian rat snake in a phylogenetic approach, none has so far led to any taxonomic changes. We assessed the intraspecific morphological variation and phylogeographic relationships among specimens from different locations across its updated distribution. Our molecular (1191 bp mtDNA, 565 bp nuDNA) and morphological data provide sufficient evidence to support three distinct lineages within the Z. hohenackeri complex with a different arrangement compared to a previous study. These represent the subspecies Z. h. hohenackeri, Z. h. tauricus, and a lineage from southwestern Turkey which is described as a new subspecies. Aspects of historical biogeography and conservation status are briefly discussed.
  • ... An ML analysis was performed using RaxML GUI v. 0.95 (Silvestro & Michalak 2012) with the 'GTRGAMMA' option and 1000 bootstrap replicates. We used the estimates of the age of the split and separation between Montivipera and Macrovipera as the calibration point, assuming this separation occurred 15.3 million years ago (Stümpel et al. 2016). ...
    Article
    A new species of blunt-nosed viper of the genus Macrovipera is described from the central and southern parts of Iran on the basis of morphological and molecular examination. The mitochondrial Cytb gene was used to investigate phylogenetic relationships amongst the Iranian species of the genus Macrovipera. A dataset with a final sequence length of 1043 nucleotides from 41 specimens from 18 geographically distant localities across Iran was generated. The findings demonstrated that two major clades with strong support can be identified within the genus Macrovipera in Iran. One clade consists of individuals belonging to a new species, which is distributed in the central and southern parts of Iran; the second clade includes two discernible subclades. The first subclade is distributed in western and northwestern Iran, Macrovipera lebetina obtusa, and the second subclade consists of northeastern populations, representing Macrovipera lebetina cernovi. The new species, Macrovipera razii sp. n., differs from its congeners by having higher numbers of ventral scales, elongated anterior chin-shields, and lower numbers of canthal plus intersupraocular scales. © 2018 Deutsche Gesellschaft für Herpetologie und Terrarienkunde e.V. (DGHT), Mannheim, Germany.
  • ... Latifi (Mertens, Darewsky & Klemmer, 1967) and M. albicornuta (Nilsen & Andren, 1985) all are belonging to M. raddei (Boettger, 1890). Another phylogenetic study conducted by Stümpel et al. (2016) suppose that M. kuhrangica and M. latifi are as valid taxa. (Mertens, Darewsky & Klemmer, 1967) Vipera latifii Mertens, Darevsky & Klemmer, 1967: 161-168. ...
    Article
    Full-text available
    During past years different studies have attempted to describe the tetrapod fauna of Iran, most of which have focused on the herpetofauna. However there is no coherent study of the endemic species of Tetrapoda in Iran. In this study, we provide a list of endemic species of Tetrapoda in Iran, mention their habitat, distribution, their conservation status (IUCN) and important biological note. Eighty endemic species of Tetrapoda occur in Iran, of which 82.50% are reptiles. Thirty-eight species (47.50% of total endemic species of Tetrapoda) have no submitted data to IUCN; of which 35 species are reptiles. Additional studies are needed to provide data about the conservation status of tetrapod fauna of Iran, especially the endemic fauna.
  • ... Ecological Niche Modelling. We compiled occurrence localities from available literature (NILSON et al., 1990;TYNIE, 1991;MULDER, 1994;1995;GÖÇMEN et al., 2009;2014, STÜMPEL et al., 2016 and our own field data. Many authors did not indicate an exact locality data in case of avoiding international pet trade. ...
    Article
    Full-text available
    The Mountain viper, Montivipera albizona, is an endemic to Anatolia and distributes in Anatolian Diagonal, Anti-Taurus Mountains and Amanos Mountains. Unfortunately, the species faced serious threats within narrow distribution range. Here, we evaluated the present status and main threats of the Mountain viper by 60-days intensive fieldwork and interviews with locals. According to our data, the Mountain viper prefers the rocky and those mountainous and rugged areas covered with less vegetation and they are active from the beginning of April to end of November. In spring, it is possible to observe 1 to 5 individuals in the suitable habitats depending on its density. We prepared 5-year Action Plan for Turkish General Directorate of Nature Conservation and National Parks and planned the roadmap for sustainability of the species with participation of regional administration, NGOs, and locals. We found that agricultural activities, overgrazing, road constructions, quarries, pet trade, sportive hunting, deliberate or accidental killing, and climatic change are the main threats on the Mountain viper in Anatolia. The main conservation measures include: creating some protected habitats in the high viper density regions, long-term monitoring survey to obtain data on its ecology and population trends, education and awareness raising activities among locals to prevent illegal collection/killing of the vipers.
  • ... Laophis crotaloides Owen, 1857, from the early Pliocene (MN 15) of the Thessaloniki area in Greece, is one of the prominent exceptions to this rule, as this gigantic (centrum length more than 15 mm) taxon is rather distinct from all other European related species and cannot be further assigned with confidence to any viperid subclade (Georgalis et al., 2016a). Anyway, the taxonomic uncertainty about the alpha taxonomy of fossil "Oriental Vipers" further hinders the understanding about the biogeography and evolutionary patterns of these large snakes, especially considering that divergence dates on the basis of molecular data between extant Macrovipera and Montivipera suggest that the two lineages had already diverged by around the middle Miocene (Langhian) (Stümpel et al., 2016). ...
    Article
    Full-text available
    We herein describe the fossil amphibians and reptiles from the Neogene (latest Miocene or earliest Pliocene; MN 13/14) locality of Maramena, in northern Greece. The herpetofauna is shown to be extremely diverse, comprising at least 30 different taxa. Amphibians include at least six urodelan (Cryptobranchidae indet., Salamandrina sp., Lissotriton sp. [Lissotriton vulgaris group], Lissotriton sp., Ommatotriton sp., and Salamandra sp.), and three anuran taxa (Latonia sp., Hyla sp., and Pelophylax sp.). Reptiles are much more speciose, being represented by two turtle (the geoemydid Mauremys aristotelica and a probable indeterminate testudinid), at least nine lizard (Agaminae indet., Lacertidae indet., ?Lacertidae indet., aff. Palaeocordylus sp., ?Scincidae indet., Anguis sp., five morphotypes of Ophisaurus, Pseudopus sp., and at least one species of Varanus), and 10 snake taxa (Scolecophidia indet., Periergophis micros gen. et sp. nov., Paraxenophis spanios gen. et sp. nov., Hierophis cf. hungaricus, another distinct “colubrine” morphotype, Natrix aff. rudabanyaensis, and another distinct species of Natrix, Naja sp., cf. Micrurus sp., and a member of the “Oriental Vipers” complex). The autapomorphic features and bizarre vertebral morphology of Periergophis micros gen. et sp. nov. and Paraxenophis spanios gen. et sp. nov. render them readily distinguishable among fossil and extant snakes. Cryptobranchids, several of the amphibian genera, scincids, Anguis, Pseudopus, and Micrurus represent totally new fossil occurrences, not only for the Greek area, but for the whole southeastern Europe. The four different types of serration within the Varanus teeth from Maramena raise questions on the taxonomic importance or the variability of this feature. The large number of distinct amphibian and reptile taxa in Maramena makes this Greek locality by far the most diverse and speciose among all European localities across the latest Miocene and earliest Pliocene. An estimation of the palaeoprecipitation value of the locality is provided. The biogeographic origins of the Maramena herpetofauna are not fully resolved, though certain of its elements were previously only known from the early and middle Miocene of Central Europe.
  • Article
    Full-text available
    Current genetic variation patterns across different geographic areas are mainly the result of various historical processes, including paleoclimate oscillations and tectonic events. Southern China, characterized by a complex paleoclimate and topography, presents an opportunity to examine such genetic variation patterns. Herein, we selected three species of moustache toads (Leptobrachium boringii, Leptobrachium liui and Leptobrachium leishanense) distributed across southern China to elucidate historical processes that shaped their current phylogeographic patterns based on two mitochondrial genes and eight microsatellite loci. L. boringii and L. liui are estimated to have genetically diverged from the late Miocene to the Pleistocene, which were largely associated with paleoclimate oscillations and geological barriers generated by complex mountain systems in southern China. Within L. liui, a recent split was caused by the rising of the Wuyi Mountains during the early Pleistocene. Demographic analyses revealed that complex topography might play buffering roles in Leptobrachium species during the last glacial maximum. These findings not only highlight the combined effects of climatic oscillations and tectonic events on driving genetic divergences, but also contribute to the general understanding of the phylogeography in this biodiversity hotspot.
  • Article
    The Anatolian Peninsula, comprising most of modern Turkey, is situated at the junction of Europe, the Middle East and Asia. Together with its complex geomorphological and climatic history, this has given rise to a rich fauna and flora, which exhibits a wide range of historical biogeographical patterns. The stone marten (Martes foina) is a small carnivore that is found across the temperate Palaearctic region including Anatolia, where it is often associated with habitats modified by humans, but few genetic data exist for this species. We sequenced a 1840-bp region of the mitochondrial genome from 97 martens sampled across the peninsula and intron 7 of the nuclear β-fibrinogen gene from 53 of these. Two mitochondrial lineages were recovered, with overlapping eastern and western distributions, but there was no geographical structure for the autosomal marker. Coalescent analyses indicated that both of the lineages originated during the Last Glacial Maximum, one of them within an eastern Anatolian refugium and the other in a western Anatolian or Balkan refugium. The western lineage colonized most of Europe in the Holocene, while the eastern lineage may be endemic to Anatolia, from where it colonized the Iberian Peninsula via human translocation. The presence of at least one refugial stone marten population highlights the importance of Anatolia to the preservation of genetic variation and biodiversity.
  • Article
    Full-text available
    The designation of taxonomic units has important implications for the understanding and conservation of biodiversity. Eurasian vipers are a monophyletic group of viperid snakes (Serpentes, Viperinae), currently comprising four genera (Daboia, Macrovipera, Montivipera and Vipera) and up to 40 species. Taxonomic units have been described using a wide variety of methods and criteria, and consequently, considerable controversy still surrounds the validity of some currently listed species. In order to promote a consensus-and evidence-based taxonomy of Eurasian vipers, we analysed published mitochondrial and nuclear DNA sequences for this group to reconstruct phylogenetic relationships among currently recognized viper species. We also compiled information on external morphology to assess their morphological distinctiveness. Phylogenetic inference based on mtDNA sequences shows contrasting levels of divergence across genera and species and identifies several instances of non-monophyly in described species. Nuclear DNA sequences show extremely low levels of genetic variation, with a widespread pattern of allele sharing among distant species, and even among genera. Revision of morphological data shows that most species designations rely on scalation traits that overlap extensively among species of the same genus. Based on our combined assessment, we recognize 15 taxa as valid species, three taxa which likely represent species complexes, 17 taxa of doubtful validity as species, and five taxa for which species status is maintained but further research is highly recommended to assess taxonomic arrangements. We stress the need to implement integrative taxonomic approaches for the recognition of evidence-based taxonomic units in Eurasian vipers.
  • Article
    Turkey hosts 5 of the 11 species of Talpa described to date and Anatolia, in particular, appears to be an important centre of diversity for this genus. Of these taxa, the Levant mole, Talpa levantis Thomas, 1906 has been found to consist of two genetically divergent sublineages, which may represent separate species. Here, we use a combination of mitochondrial and nuclear DNA sequences from specimens of T. levantis s.lat., collected across the species’ geographical range, to explore the systematics and demographic history of Levant moles. Both mitochondrial and nuclear markers confirm the existence of distinct eastern and western sublineages, which apparently diverged from each other in the early Pleistocene. Given the degree of cytochrome-b divergence between these (7.28%), we consider them to represent independent cryptic species. By including topotypic specimens of T. levantis s. str. in our study, we are able to show that this name applies to the western sublineage, distributed across most of the Anatolian Black Sea coastal region, from the vicinity of Trabzon in the east, westwards to Marmara. The earliest name available for the eastern taxon, found in Transcaucasia and adjacent parts of north-eastern Anatolia, is T. transcaucasica Dahl, 1944. Cytochrome-b haplotype diversity in T. levantis is relatively high and demographic analyses suggest that the species may have survived in multiple, separate, refugial areas during the Pleistocene. Our work brings the total number of named mole species recognized in Turkey to six, emphasising the importance of this region as a global centre of mole diversification.
  • Article
    Episodes of population growth and decline leave characteristic signatures in the distribution of nucleotide (or restriction) site differences between pairs of individuals. These signatures appear in histograms showing the relative frequencies of pairs of individuals who differ by i sites, where i = 0, 1, .... In this distribution an episode of growth generates a wave that travels to the right, traversing 1 unit of the horizontal axis in each 1/2u generations, where u is the mutation rate. The smaller the initial population, the steeper will be the leading face of the wave. The larger the increase in population size, the smaller will be the distribution's vertical intercept. The implications of continued exponential growth are indistinguishable from those of a sudden burst of population growth Bottlenecks in population size also generate waves similar to those produced by a sudden expansion, but with elevated uppertail probabilities. Reductions in population size initially generate L-shaped distributions with high probability of identity, but these converge rapidly to a new equilibrium. In equilibrium populations the theoretical curves are free of waves. However, computer simulations of such populations generate empirical distributions with many peaks and little resemblance to the theory. On the other hand, agreement is better in the transient (nonequilibrium) case, where simulated empirical distributions typically exhibit waves very similar to those predicted by theory. Thus, waves in empirical distributions may be rich in information about the history of population dynamics.
  • Article
    Full-text available
    Arlequin ver 3.0 is a software package integrating several basic and advanced methods for population genetics data analysis, like the computation of standard genetic diversity indices, the estimation of allele and haplotype frequencies, tests of departure from linkage equilibrium, departure from selective neutrality and demographic equilibrium, estimation or parameters from past population expansions, and thorough analyses of population subdivision under the AMOVA framework. Arlequin 3 introduces a completely new graphical interface written in C++, a more robust semantic analysis of input files, and two new methods: a Bayesian estimation of gametic phase from multi-locus genotypes, and an estimation of the parameters of an instantaneous spatial expansion from DNA sequence polymorphism. Arlequin can handle several data types like DNA sequences, microsatellite data, or standard multilocus genotypes. A Windows version of the software is freely available on http://cmpg.unibe.ch/software/arlequin3.
  • Article
    Lizards, snakes and amphisbacnians together constitute the Squamata, the largest and most diverse group of living reptiles. Despite their current success, the early squamate fossil record is extremely patchy. The last major survey of squamate palaeontology and evolution was published 20 years ago. Since then, there have been major changes in systematic theory and methodology, as well as a steady trickle of new fossil finds. This review examines our current understanding of the first 150 million years of squamate evolution in the light of the new data and changing ideas. Contrary to previous reports, no squamate fossils are currently documented before the Jurassic. Nonetheless, indirect evidence predicts that squamates had evolved by at least the middle Triassic, and had diversified into existing major lineages before the end of this period. There is thus a major gap in the squamate record at a time when key morphological features were evolving. With the exception of fragmentary remains from Africa and India, Jurassic squamates are known only from localities in northern continents (Laurasia). The situation improves in the Early Cretaceous, but the southern (Gondwarian) record remains extremely poor. This constrains palaeobiogeographic discussion and makes it difficult to predict centres of origin for major squamate clades on the basis of fossil evidence alone. Preliminary mapping of morphological characters onto a consensus tree demonstrates stages in the sequence of acquisition for some characters of the skull and postcranial skeleton, but many crucial stages - most notably those relating to the acquisition of squamate skull kinesis - remain unclear.
  • Article
    Prangos abieticola Aytaç & H.Duman (Apiaceae) from southern Anatolia, Turkey is described and illustrated. This new taxon grows on calcareous rocks in open Abies cilicica (Antoine & Kotschy) Carrière and Pinus nigra J.F.Arnold subsp. nigra var. caramanica (Loudon) Rehder forest in the middle of the Taurus Mountains (C3 Antalya and Konya Province). It is endemic and closely related to Prangos heyniae H.Duman & M.F.Watson and P. ferulacea (L.) Lindl. The diagnostic morphological characters of similar taxa are discussed.
  • Article
    Up to now, the oldest known colubrid snakes were fossils from the early Oligocene of Western Europe and from the Arabian Peninsula. An indeterminate colubrid has been recently recovered in the late Eocene of Thailand. The presence, in Asia, of the oldest known colubrid snake is consistent with an origin of this family in this continent. -Authors
  • Article
    This contribution spotlights the present research on the (oreal) subalpine to subnival mountain vegetation of Turkey. It concisely compiles our present knowledge, which is very heterogeneous in geographical terms. The situation is comparatively good in NW and W Anatolia and the western half of the Taurus range, from where a first consolidation stage can be reported. By contrast, the E Taurus remains a largely unexplored area. The North Anatolian chains have hitherto attracted a few local researchers only, and those important pioneering accounts cover less than half of the vegetation types actually present. The results of the last years' field work in the Taurus range represent the core of the presentation and focus on the syntaxonomy and phytogeography of the Tauric System. An annotated conspectus of all known major syntaxa (alliances to class group level) recorded from the Anatolian mountains is presented. It includes 15 first records of mostly Euro-Siberian and Mediterranean orders and classes that proved to be incorrectly interpreted, completely unstudied or as yet unreported. The classification of the N Anatolian mountain vegetation suffered severely by neglecting the rich literature about the neighbouring mountain chains of the Balkans or the Great and Little Caucasus. In taking a broader view, some major vegetation units are re-defined to better reflect Turkey's position in the Eurasian Alpic-Himalayan fold mountain system. A phytogeographical subdivision of the Turkish mountain ranges and all of the Tauric System based on both the asylvatic high mountain vegetation and flora is provided. The subdivision is fully backed by the pattern revealed by the forest communities. Despite the achievements in the past, such a lecture is to a certain extent a tale of ignorance. Pointing out the gaps in our knowledge may help to co-ordinate the studies needed. Another goal is to discuss conflicting or even inappropriate syntaxonomic concepts and working methods to come closer to a common base for future vegetation surveying projects.
  • Article
    Full-text available
    Prangos turcica A. Duran, M. Saǧiroǧlu & H. Duman sp. nova (Apiaceae) is described and illustrated from Anatolia, Turkey. The species grows in mixed forest, on shady slopes and open forest in Amanos Mountains (C6 Osmaniye province). Prangos turcica, an endemic confined to South Anatolia, is closely related to P. trifida (Mill.) Herrnst. & Heyn. Diagnostic morphological characters from closely similar taxa are discussed. Habitat ecology of the species is also discussed. Prangos turcica is diploid with the chromosome number of 22.
  • Article
    Full-text available
    North-eastern Anatolia harbours a high diversity of viperid snakes with only a limited knowledge about their distribution and with relationships among these vipers not yet fully resolved. Moreover, information on habitat attributes for most of these vipers is scarce. We initiated a multi-year project to improve our knowledge on their distribution and habitat preferences, especially by searching contact zones of closely related and ecologically similar species and evaluate potential gene flow and species integrity. In this context and as an intermittent step, we report new localities nearby putative contact zones. Thus, herein we present new information on the distribution of Vipera barani, V. kaznakovi, V. darevskii, V. eriwanensis, Montivipera wagneri, M. raddei and Macrovipera lebetina based on our field work and third sources provided to us. With these data, we were able to reduce the distribution gaps between three pairs of "parapatric", related or ecologically similar, viper species (genus Vipera) by mostly 50%, and detected a putative contact zone in a fourth species pair (genus Montivipera). All putative contact zones are discussed in an ecological context. In addition, we add new sites of M. lebetina in the Province Artvin and discuss its northern limit in Turkey.
  • Article
    The true global incidence of envenomations and their severity remain largely misunderstood, except for a few countries where these accidents are rare or are correctly reported. Nevertheless, this information is essential for drawing up guidelines for dealing with snake-bites, to plan drug supplies, particularly antivenin, and to train medical staff on snake-bite treatments. Since the comprehensive review by Swaroop & Grab in 1954 no global survey has been carried out on snake-bite epidemiology. The present article is an attempt to draw the attention of health authorities to snake envenomations and urges them to prepare therapeutic protocols adapted to their needs.
  • Article
    Paleogeographical considerations on the development of the Paratethys and the Mediterranean during Oligocene and Miocene are presented in twelve time-slices. Plate tectonic activities and the collision of India with Asia caused the destruction of the Western Tethys Ocean in the Late Eocene. The Mediterranean and the intracontinental Paratethys basins came into existence as new marine realms. In the Mediterranean Basin open oceanic connections existed throughout the Oligocene and most of the Miocene. The Eastern Paratethys and the Central to Western Paratethys showed different marine conditions and changing connections most of the time. A first period with reduced salinity, anoxic bottom conditions, and strong endemisms occurred throughout the Paratethys in a short period of the Lower Oligocene (Solenovian, NP 23). It was followed by more open marine conditions with wide-spread clastic sedimentation (Upper Kiscellian, Kalmykian, NP 24). By the collision of Africa and Arabia with Eurasia, the seaway between the Mediterranean Sea and the Indian Ocean was closed in Burdigalian time, but a new landbridge enabled a distinct mammal migration between the continents (Gomphotherium Landbridge). During the Middle Miocene marine seaways between the Indian Ocean, the Mediterranean, and the Paratethys opened and closed intermittently. Finally, the marine connections of the Paratethys were strongly reduced, and gave way to the endemic faunal development during the later Miocene (Sarmatian to Pontian).
  • Article
    Vipera latifii Mertens, Darevsky & Klemmer, a recently described viper from northern Iran, is in severe need of conservation. Its range is restricted to the unique, alpine Lar Valley, which in a few years will be used as a huge water reservoir. Observations on the biology of Vipera latifii are given. Sympatric amphibians and reptiles show ecotypic adaptations.
  • Article
    Full-text available
    The Mio-Pliocene basins around the Antalya gulf in SW Turkey developed above the Tauric Mesozoic platforms on which the Antalya nappes had been thrusted (in Late Cretaceous-Paleocene times). The closure of the initial Isparta Angle during these events (E-W compression) initiated the N-S orientation of the main structural lines, which persisted later and explains the orientation of the Aksu basin in contrast with the E-W orientation of the eastern Neo-gene Mediterranean basins. The area, and all southwestern Turkey, became emergent at the end of the Oligocene and were the site of shallow-marine carbonate deposits in the Chattian-Aquitanian, giving way to the wide Lycian basin in Burdigalian-Langhian times. The progressive emplacement of the Lycian nappes from the north over this basin provoked first its subsidence and then its emersion when the nappes attained their final position over the Bey Daglari platform in Langhian times. Coinciding, or in response to the Lycian nappes emplacement, the Aksu basin was initiated as an elongated N-S graben which was filled by thick accumulations of terrestrial and marine deposits(including coral reefs), which derived from the erosion of the Lycian allochton and its basement (Langhian?, Serravallian and Tortonian times). The syn-sedimentary tectonics : reactivation of the normal faults along the west margin of the basin, the continuous uplift of the neighbouring continental areas (beginning of the Aksu thrust), governed the geometry of the basin. As a result and due to the uplift of its northern margin, the Aksu basin migrated towards the south and in Messinian times it was reduced to a narrow gulf along the eastern margin of which the Gebiz limestones were deposited as fringing coral reefs. The age of these limestones has been debated. Our new data allow us to attribute them to the Messinian. The drastic retreat of the sea at the end of this period, provoked the erosion of large parts of the Messinian deposits and the formation of deep canyons on land and under the sea down to the Antalya abyssal plain, in which evaporites were deposited. During the Zanclean transgression, the Eskiköy-Kargi canyon was filled by coarse clastics of a Gilbert delta derived from the northern continental area following a model well known elsewhere in the Mediterranean basins. Southward , shallow-marine sands and marls unconformably cover the remnants of the Messinian deposits and the emergent areas of the southern Antalya gulf. After Zanclean times (end of Pliocene?), the Aksu basin was deformed, due to the west-directed Aksu compressional event (end of the Aksu thrust). Quaternary terraces of the Aksu river at various altitudes, as well as the terraces of the Antalya tufa can be related to sea level fluctuations. Evolution fini-tertiaire du bassin de l'Aksu au sud-ouest de la Turquie. Nouvelles données Mots-clés. – Méditerranée orientale, Sud-ouest de la Turquie, Golfe d'Antalya, Bassins néogènes et quaternaires, Tectonique syn-sé-dimentaire et eustatisme, Crise de salinité messinienne. Résumé. – Les bassins néogènes du golfe d'Antalya (d'ouest en est : Aksu, Köprü, Manavgat) se forment sur un substra-tum composite incluant les plates-formes tauriques (Bey Daglari à l'ouest et Anatolie à l'est) surmontées par les nappes d'Antalya issues du Bassin pamphylien (Néotéthys sud) au cours du Crétacé supérieur-Paléocène, et leur couverture pa-léogène. La convergence E-W et la collision des plates-formes induisent des structures orientées N-S. Cette direction persistera ensuite et explique l'orientation actuelle du bassin de l'Aksu contrastant avec l'orientation E-W des bassins est-méditerranéens. Cet ensemble est recouvert au Chattien-Aquitanien par une mer peu profonde à dépôts carbonatés récifaux et péri-récifaux : la formation de Karabayir qui couvrait tout le SW de la Turquie. Sous l'effet de l'avancée des nappes lyciennes (depuis le nord), au Burdigalien et au Langhien inférieur, cette plate-forme devient subsidente au NW et le bassin lycien se creuse et devient profond. Il est alimenté en détritiques terrigènes à partir du front des nappes ly-ciennes depuis le NW. La fin du Langhien est marquée par le développement de récifs de coraux dans un contexte de grands épandages deltaïques. C'est à cette époque que se forme véritablement le bassin d'Antalya, en coincidence, ou en conséquence de la mise en place des nappes lyciennes : l'ensemble du Taurus émerge, sauf le bassin d'Antalya qui s'effondre le long de failles N-S. En bordure est, la faille de Kirkavak est active. En bordure ouest, le relief abrupt à la limite des Bey Daglari est interprété comme une faille. On a donc deux mouvements en sens inverse : soulèvement des plates-formes et effondrement du bassin. Ce dernier est alors fortement réduit en surface par rapport au bassin lycien. Manuscrit déposé le 21 janvier 2009 ; accepté après révision le 12 juillet 2010. Le Serravallien est mal connu. Le Tortonien est représenté par d'énormes masses de conglomérats fluvio-marins dans lesquels s'intercalent des récifs de coraux, le long des bordures, passant à des turbidites marno-gréseuses dans les parties un peu plus profondes du bassin. Le bassin de l'Aksu est alors bien individualisé par rapport aux autres bassins du golfe d'Antalya. Le Messinien n'est connu que ponctuellement et on ne peut pas reconstituer complètement le bassin à cette période. Il devait avoir une géométrie voisine de celle du bassin Pliocène, c'est-à-dire qu'il était réduit par rapport au bassin tortonien et emboîté dans ce dernier au sud. Le faciès messinien le plus caractéristique est représenté par les calcaires de Gebiz (coraux et calcaires bioclastiques). Au nord du bassin, ils sont discordants sur le socle et plus au sud sur le Tortonien. Intercalés entre du Messinien (NN11b) et du Pliocène basal (NN12), on peut donc leur attribuer un âge Messinien. Entre les calcaires de Gebiz et le Pliocène une discontinuité résulte selon toute vraisemblance de la crise de salinité messinienne (présence également de blocs de gypse). Une érosion a accompagné la baisse du niveau marin lors de la crise de salinité comme en témoigne un canyon à parois abruptes creusé dans le Tortonien. Au Pliocène inférieur, la mer réinvestit le bassin et les canyons alors que la moitié nord du bassin émerge et le bassin pliocène s'encastre dans le bassin tortonien-messinien. Le bassin de l'Aksu se réduit, pendant que ses bordures continuent à se soulever. A la fin du Miocène et au moins jusqu'après le Zancléen, une compression d'est en ouest provoque des écaillages profonds dans le bassin et l'empilement d'écailles vers l'ouest. Cet événement (charriage de l'Aksu) affecte toute la série mio-plio-cène. Au-dessus du chevauchement principal, le Tortonien peut être porté à près de 2 500 m d'altitude. Jusqu'au Plio-cène la tectonique joue donc un rôle très important dans le bassin de l'Aksu. C'est au Quaternaire que les mouvements eustatiques sont les plus évidents. Les travertins d'Antalya (ou tufa) se sont déposés à l'air libre dans des bassins ali-mentés par des résurgences. La terrasse supérieure est à + 300 m pendant que la terrasse la plus basse est à – 40 m. Le fleuve Aksu a déposé des terrasses alluviales anciennes dont les plus hautes sont à + 70 m pendant que la base de son lit récent est à –138m. Au niveau du barrage de Kargi, l'Aksu a creusé par surimposition dans les calcaires massifs méso-zoïques, une gorge très étroite de 200 m de profondeur, juste à côté du canyon messinien (rempli de Pliocène). Cette gorge permet à la rivière de franchir l'abrupt qui sépare le basin tortonien du bassin messinien-pliocène. Le creusement est post-Pliocène, sans doute en rapport avec les mouvements eustatiques du Quaternaire. En mer, il existe également un grand canyon qui ne se trouve pas dans le prolongement de l'Aksu actuel mais qui est décalé vers l'ouest, et localisé près de la bordure ouest du golfe d'Antalya. Il est probable qu'il résulte du creusement messinien de l'ancien cours du fleuve Aksu. L'ampleur des mouvements verticaux est donc très importante (au moins 4000 m pour le Miocène inférieur). L'emboîtement des bassins plus récents dans des bassins plus anciens, se produit successivement au Miocène moyen, au Messinien-Pliocène et au Quaternaire. A chaque étape le continent se soulève pendant que les bassins s'effondrent. Le rôle de la tectonique est évident mais de quelle tectonique s'agit-il ? Le contexte du bassin d'Antalya est en effet parti-culièrement compliqué du fait de l'interférence : 1– d'une tectonique active en extension E-W à NE-SW avec des failles N-S et NW-SE; 2– d'une compression de l'est vers l'ouest et 3– d'une subduction de la Méditerranée du sud vers le nord sous la ride Florence et sous le bassin d'Antalya (et jusque sous le Taurus ?). Bien que pas forcément compatibles entre eux ces trois types de tectonique sont actifs ainsi que le suggère la sismicité régionale.
  • Article
    Population structure is the result of both present processes and past history. Molecular markers are proving of great value in describing the former, and it is important to similarly determine the latter in order to understand their respective contributions. The study of palaeo-climates has also advanced significantly, and in particular that of the Pleistocene ice ages, which modified species ranges considerably. The last ice age and rapid post-glacial colonization of Europe is summarized. Possible population genetic consequences of expansion northward from southern refugia, and those of remaining in these mountainous regions are discussed. A series of recent case studies are detailed where DNA sequence information has been used to describe species genetic variation and subdivision across Europe. These include a grasshopper, the hedgehog, oak trees, the common beech, the black alder, the brown bear, newts, shrews, water vole, silver fir and house mice. These molecular data confirm southern peninsulas of Europe as major ice age refugia, and in most cases demonstrate that genetically distinct taxa emerged from them. They can thus define genomic differences and so greatly augment previous fossil data. The refugial genomes contributed differently in various species to the re-colonization of Europe, with three broad patterns described as paradigms—«grasshopper», «hedgehog» and «bear». These different expansion patterns produced clusters of hybrid zones where they made contact, and it is argued that many species genomes may be further cryptically subdivided. A reduction in diversity from southern to northern Europe in the extent of allelic variation and species subdivision is seen; this is attributed to rapid expansion northward and the varied topography of southern refugia allowing populations to diverge through several ice ages. The differences in DNA sequence indicate that some species have been diverging in refugial regions for a few ice ages at most, whilst distinct lineages in other species suggest much more ancient separation.
  • Article
    Full-text available
    Theory predicts low niche differentiation between species over evolutionary time scales, but little empirical evidence is available. Reciprocal geographic predictions based on ecological niche models of sister taxon pairs of birds, mammals, and butterflies in southern Mexico indicate niche conservatism over several million years of independent evolution (between putative sister taxon pairs) but little conservatism at the level of families. Niche conservatism over such time scales indicates that speciation takes place in geographic, not ecological, dimensions and that ecological differences evolve later.
  • Book
    Phylogeography of Southern European Refugia provides the first synthesis of the remarkable diversity, evolutionary complexity, and conservation importance of the flora and fauna in the Mediterranean region, with emphasis on the three major peninsular refugia. The book frames a number of case studies within several introductory and closing chapters that review the development and importance of phylogeographic approaches in evolutionary biology. The book opens with a chapter by the field’s founding father, John Avise, reviewing the 25 evolutionary insights that have arisen from the phylogeographic revolution. Key to this perspective is the notion that the multidisciplinary field of phylogeography is serving as an epistemological bridge between the formerly distinct fields of population genetics and phylogenetics. Next, the foremost leaders of plant phylogeography in Europe, Rémy Petit and Giovanni Vendramin provide an extraordinarily comprehensive and studious review of the history of plant organelle genetics and their application in phylogeography. A closing chapter provides an up-to-date review of the caveats, challenges, and future prospects in phylogeography as a discipline. The volume contains both broad scale and regional phylogeographic studies with original data from a range of organisms including trees, mammals, insects, fish, reptiles, and amphibians. Throughout the volume it is clear that phylogeographic studies, especially in the major peninsulas of southern Europe (Iberia, Italy, the Balkans) have provided enlightening insights into the evolutionary complexity and conservation importance of biota occupying Europe’s most prominent glacial refugia
  • Article
    Recent studies of the reptile fauna in central Zagros mountains have concluded in the finding of a single specimen of a new taxon in the raddei species group in the genus Montivipera. In addition a photo of a second specimen from the same locality — the Kuhrang region, Chaharmahal and Bakhtiari Province, Iran is available. Although there is only a single specimen at hand we are not able to compare the variation in morphological features with known taxa of Montivipera raddei species group, but the morphological characteristics of this single specimen separates it from any former known taxa within Montivipera. It clearly belongs to the M. raddei species group and in some features it is similar to M. latifii but unique in color patter and other characters. We consider it to be a sep-arate and isolated new species, Montivipera kuhrangica. The species seems to be very rare and is certainly highly exposed to a risk of habitat destruction and it needs conservation programs.
  • Article
    Full-text available
    We report new localities and range extensions for four vipers from Turkey. The distribution of Montivipera wagneri is extended substantially southwards into Muş province, showing that this species is not endemic to the Aras valley as previously stated in the literature. Similarly, an unverified photograph of Montivipera albizona from Erzincan province, at its currently known eastern limit, was confirmed with new material, and its range is extended substantially southwards to the Anamos (Nur) Mountains in Hatay province. Smaller range extensions are reported for Vipera ammodytes transcaucasiana and V. darevskii.
  • Article
    Full-text available
    New data on Syrian amphibians and reptiles collected between 1990 and 2004 are given. The most important findings are the Elap~d snake Walterznnesza aegyptza, here reported for the first time from Syna; Zamenzs hohenackerz and Vzpera xanthzna, previously known only from Mount Hermon; and the very rare and poorly known Telescopus nzgriceps and Lytorhynchus ken-nedyz. Other specles were previously known from only a very few Syrian localities. Kurzfassung. Es werden neue Daten iiber syrische Arnphlbien und Reptilien mitgeteilt, dle nvischen 1990 und 2004 gesammelt wurden. Die wichigsten Funde stellen Walterinnesza aegyp-tia dar, d ~ e hier mm ersten Mal aus Synen gemeldet wird; Zamenis hohenackeri und Vzpera xan-thma, die blsher nur vom Mount Hermon bekannt waren, und die sehr seltenen und wemg be-kannten Arten Telescopus nigrzceps und Lytorhynchus kennedyi. Fiir andere Arten waren bisher OR nur Einzelnachweise bekannt.
  • Article
    This is the first comprehensive book on the weather and climate of mountains. The first chapter on the historical aspects is followed by chapters dealing with the geographical controls of mountain meteorological elements, circulation systems related to orography and the climatic characteristics of mountains. Chapter 5 looks at case studies and Chapter 6 bioclimatology. The final chapter examines the evidence for and the significance of changes in mountain climates.-D.G.Tout
  • Article
    Full-text available
    The distribution of Tettigoniinae (Orthoptera, Tettigoniidae) specieswithin four phytogeographical provinces in Turkey is reviewed and presented onthe basis of field and collection studies during 1987–2001 andmiscellaneous data from previous literature. One hundred and sixty speciesbelonging to 28 genera have been recorded. The Anatolian species constituteapproximately 30% of the world-wide species of the subfamily, with a very highproportion (=83%) being endemic or semi-endemic, nearly all of which arebrachypterous. Regarding the numbers of total and endemic species, theprovinces can be ordered as follows: Mediterranean > Irano-Anatolian >Euxin > Mesopotamia. Also, Anatolian Tettigoniinae have a very high rate ofendemism in each province; 84.1% in the Mediterranean, 78.1% in Irano-Anatolia,56.3% in Mesopotamia and 42.4% in Euxin. The two provinces in Anatolia havingthe greatest species diversity are also the two most mountainous provinces, soit is suggested that a primary factor in Tettigoniinae diversity is theAnatolian Taurus mountains. Finally, some conclusions are drawn on biodiversityand conservation of Tettigoniinae species in Anatolia and I have suggested 23species to be included in the 'IUCN Red List' under VU B2+ac.
  • Article
    Aim The aim of this study was to elucidate the phylogeographical pattern of taxa composing the Vipera ursinii complex, for which the taxonomic status and the dating of splitting events have been the subject of much debate. The objectives were to delimit potential refugia and to date splitting events in order to suggest a scenario that explains the diversification of this species complex. Location Western Europe to Central Asia. Methods Sequences of the mitochondrial cytochrome b and NADH dehydrogenase subunit 4 (ND4) genes were analysed for 125 individuals from 46 locations throughout the distribution range of the complex. The phylogeographical structure was investigated using Bayesian and maximum likelihood methods. Molecular dating was performed using three calibration points to estimate the timing of diversification. Results Eighty-nine haplotypes were observed from the concatenation of the two genes. Phylogenetic inferences supported two main groups, referred to in this study as the ‘ursinii clade’ and the ‘renardi clade’, within which several subclades were identified. Samples from Greece (Vipera ursinii graeca) represented the first split within the V. ursinii complex. In addition, three main periods of diversification were revealed, mainly during the Pleistocene (2.4–2.0 Ma, 1.4 Ma and 1.0–0.6 Ma). Main conclusions The present distribution of the V. ursinii complex seems to have been shaped by Quaternary climatic fluctuations, and the Balkan, Caucasus and Carpathian regions are identified in this study as probable refugia. Our results support a south–north pattern of colonization, in contrast to the north–south colonization previously proposed for this complex. The biogeographical history of the V. ursinii complex corroborates other biogeographical studies that have revealed an east–west disjunction (situated near the Black Sea) within a species complex distributed throughout the Palaearctic region.