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1.
    
We provide a review and synthesis of key findings in phylogeographic research on terrestrial animals in the Aegean archipelago (Greece) and surrounding regions (Greek mainland, southern Balkans and Asia Minor). A critical review of more than 100 phylogeographic articles on 76 animal genera (30 invertebrates and 46 vertebrates) that have been published so far for the region leads to the recognition of three types of distribution patterns: old ‘colonizers’ diversified in the Aegean before the formation of the mid‐Aegean Trench (MAT – i.e. before 9 Mya), post‐MAT colonizers that arrived to the Aegean in late Miocene and Pliocene and new colonizers that inhabited the region in the Pleistocene or even the Holocene. Several problems, mainly regarding the use of calibration points and/or molecular clock rates for clade chronology, have been identified in many analyses. Despite the large amount of phylogeographic work concerning the Aegean and surrounding regions, many groups remain unstudied (especially invertebrates, micro‐organisms, fungi and plants), and many issues still remain unresolved. The relative roles of extinction, speciation, dispersal and vicariance, as well as the effects of adaptive and non‐adaptive components of diversification, need further analyses with modern tools that can provide deeper insights. A more detailed reconstruction of the palaeogeography of the region is also of prime importance. The critical views presented herein may prove useful also for the evaluation of similar work in other regions.  相似文献   

2.
    
The desiccation of the Mediterranean Sea during the Messinian Salinity Crisis 6.0-5.3 million years ago (Ma), caused a major extinction of the marine ichthyofauna of the Mediterranean. This was followed by an abrupt replenishment of the Mediterranean from the Atlantic after the opening of the Strait of Gibraltar. In this study, we combined demographic and phylogeographic approaches using mitochondrial and nuclear DNA markers to test the alternative hypotheses of where (Atlantic or Mediterranean) and when (before or after the Messinian Salinity Crisis) speciation occurred in the Mediterranean damselfish, Chromis chromis. The closely related geminate transisthmian pair Chromis multilineata and Chromis atrilobata was used as a way of obtaining an internally calibrated molecular clock. We estimated C. chromis speciation timing both by determining the time of divergence between C. chromis and its Atlantic sister species Chromis limbata (0.93-3.26 Ma depending on the molecular marker used, e.g. 1.23-1.39 Ma for the control region), and by determining the time of coalescence for C. chromis based on mitochondrial control region sequences (0.14-0.21 Ma). The time of speciation of C. chromis was always posterior to the replenishment of the Mediterranean basin, after the Messinian Salinity Crisis. Within the Mediterranean, C. chromis population structure and demographic characteristics revealed a genetic break at the Peloponnese, Greece, with directional and eastbound gene flow between western and eastern groups. The eastern group was found to be more recent and with a faster growing population (coalescent time = 0.09-0.13 Ma, growth = 485.3) than the western group (coalescent time = 0.13-0.20 Ma, growth = 325.6). Our data thus suggested a western origin of C. chromis, most likely within the Mediterranean. Low sea water levels during the glacial periods, the hydrographic regime of the Mediterranean and dispersal restriction during the short pelagic larval phase of C. chromis (18-19 days) have probably played an important role in C. chromis historical colonization.  相似文献   

3.
During the Late Miocene the Mediterranean experienced a period of extreme salinity fluctuations known as the Messinian Salinity Crisis (MSC). The causes of these high amplitude changes in salinity are not fully understood but are thought to be the result of restriction of flow between the Mediterranean and Atlantic, eustatic sea level change and climate. Results from a new Atmospheric General Circulation Model (AGCM) simulation of Late Miocene climate for the Mediterranean and adjacent regions are presented here. The model, HadAM3, was forced by a Late Miocene global palaeogeography, higher CO2 concentrations and prescribed sea surface temperatures. The results show that fluvial freshwater fluxes to the Mediterranean in the Late Miocene were around 3 times greater than for the present day. Most of this water was derived from North African rivers, which fed the Eastern Mediterranean. This increase in runoff arises from a northward shift in the intertropical convergence zone caused by a reduced latitudinal gradient in global sea surface temperatures. The northwards drainage of the Late Miocene Chad Basin also contributes. Numerical models designed to explore Late Miocene salt precipitation regimes in the Mediterranean, which typically make use of river discharge fluxes within a few tens of percent of present-day values, may therefore be grossly underestimating these fluxes.Although the AGCM simulated Late Miocene river discharge is high, the model predicts a smaller net hydrologic budget (river discharge plus precipitation minus evaporation) than for present day. We discuss a possible mechanism by which this change in the hydrologic budget, coupled with a reduced connection between the Mediterranean and the global ocean, could cause the salinity fluctuations of the MSC.  相似文献   

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Aim To infer the phylogenetic relationships and biogeography of Hydromantes, with special emphasis on the European taxa. In particular, we aimed to test: (1) the monophyly of the European species and current views on their interrelationships; and (2) previously proposed timings of the separation of European and American Hydromantes, and of biogeographically important events within Europe. Location California and the Western Mediterranean Basin, specifically south‐east France, Italy, and the island of Sardinia. Methods Partial sequences of mitochondrial genes (cytochrome b and 12S rRNA) were obtained from 45 specimens of Hydromantes, including all European extant species and subspecies, and two species from California. In addition, a fragment of the mitochondrial 16S rRNA gene was amplified for 16 specimens. Data sets were aligned using Clustal X, and well‐supported phylogenetic trees were produced using maximum‐likelihood, Bayesian and maximum‐parsimony methods. Estimates of divergence times were obtained with the program r8s , the molecular clock being calibrated using the opening of the Strait of Gibraltar, the final event in the Messinian Salinity Crisis of 5.3 Ma. Results Separation between the American and European clades occurred approximately 13.5 Ma, most probably before or after westward dispersal across the Bering Land Bridge. In Europe, divergence started in the late Miocene, when Hydromantes (A.) genei separated from other members of the genus 9 Ma and colonized south‐west Sardinia. Movement between the European mainland and Sardinia, by a member of the subgenus Speleomantes, occurred in the Messinian Salinity Crisis, after the Mediterranean Basin desiccated almost completely 5.96 Ma. Subsequent widespread aridification fragmented the geographical ranges of Hydromantes, which live in cool and humid conditions, resulting in the origin of the six species in the subgenus Speleomantes. In contrast, a second period of diversification, in continental Europe 2–1.3 Ma, was probably caused by very cold interludes during the climatic oscillations that characterized the Pleistocene. Main conclusions The molecular clock used here indicates that the separation of Californian and European Hydromantes occurred more recently than previously believed, and the same is true of some subsequent phylogenetic divergences within Europe. Estimated dates for these divergence events are consistent with known geophysical and climatic events that could have caused or facilitated them.  相似文献   

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Integrated data of calcareous plankton and benthic foraminifers from the pre-evaporitic interval of Trave section (Central Italy) allowed the reconstruction of surface and bottom-water conditions in the Central Mediterranean during the interval from 7.61 to 6.33 Ma, preceding the Messinian Salinity Crisis.Our data point out a three-step paleoenvironmental evolution. During the first stage (7.61-7.02 Ma) benthic foraminiferal assemblages depict stable, well-oxygenated and ventilated bottom-water conditions, while the surface water records variable temperature and high nutrient conditions, probably associated with strong seasonality. The second stage (7.02-6.70 Ma) points to unfavourable bottom-water condition, triggered by deep-sea stagnation. This is witnessed by a significant decrease in oxygen concentration and biotic diversity, and by the presence of stress-tolerant taxa. A general warming of the surface water and a strongly stratified water column, characterized by an expanded mixed layer, are also recorded.From 6.70 Ma onwards (third stage), a prominent change to more restricted, low-oxygenated, hypersaline conditions at the sea floor is testified by the total disappearance of deep-dwelling planktonic foraminifers and the increasing abundance of stress-tolerant species. Calcareous plankton reflects high instability of the surface water in terms of nutrients, temperature and salinity. During this stage the environmental deterioration reaches intermediate depths in the water column.The initial change toward a step-wise isolation of the Central Mediterranean bottom-waters is probably related to a general warming, responsible for a first slowing-down of the vertical circulation, favouring stratification of surface and intermediate waters and stagnation of bottom-waters. This warming is related to the restricted connection between the Mediterranean Sea and the Atlantic Ocean, which occurred since 7.146 Ma.In the Trave section, the isolation of bottom-waters most likely occurred at the same time as in other Mediterranean sections. However, due to the presence of a hiatus it cannot be excluded that it occurred with a delay of ~ 100 kyr, probably related to the shallower paleodepth of the basin.  相似文献   

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Aim To test the importance of the Lago Mare stage of the Messinian Salinity Crisis for the dispersal and diversification of European leuciscins (Cyprinidae: Leuciscinae). Location Europe. Methods Cytochrome b sequences of European leuciscins were employed to investigate phylogenetic relationships among species, using Bayesian inference, and to estimate times of diversification, using a relaxed molecular clock. The distributions of 190 European leuciscins were compiled, and regional species compositions were compared using a taxonomic similarity index and an area cladogram. Results Leuciscins restricted to the Iberian and Italian peninsulas and the West and South Balkan regions are phylogenetically more closely related to northern European species than to species from another southern European area. Application of a relaxed molecular clock to a Bayesian phylogeny indicates that most southern clades originated and diversified prior to the Messinian. Southern European regions are taxonomically distinct from one another, and from a more taxonomically homogeneous group of areas that includes Anatolia, East Balkans, Middle East, North Europe and West Russia. Main conclusions The scenario of a Messinian period of dispersal of Paratethyan fauna into Mediterranean regions, via the Lago Mare, predicts a rapid period of diversification and a pattern of close association among southern European faunas. Phylogenetic relationships among leuciscins, the timing of cladogenic events, and the taxonomic similarity among geographical regions do not conform to this expectation. The depth of clades endemic to southern Europe, together with the high levels of endemism in these regions, suggests that the faunas in these regions diverged prior to the Messinian and have evolved largely in isolation from one another. Our results support a model of gradual colonization of Mediterranean regions since the Oligocene. Subsequent connections between adjacent areas may have occurred in the Messinian or Pleistocene.  相似文献   

8.
The Fortuna Basin is an example of a marginal Mediterranean basin with evaporitic sedimentation during the Late Tortonian and Messinian. This basin shows an early restriction event before the main Messinian Salinity Crisis (MSC) that allows the Tortonian Salinity Crisis (TSC) to be proposed as a tectonic uplift event isolating the eastern Betic basins. Four evaporitic events are present in the central part of the Fortuna Basin, from bottom to top: Los Baños Marls Formation (composed by Fenazar Conglomerate Bed, Lower Gypsum Member [Mb] and Sanel Mb), Tale Gypsum Formation (Fm), Chicamo Diatomites and Gypsum Cycles Fm, and Rambla Salada Gypsum Fm. The present work documents the first biostratigraphic dating based on calcareous nannoplankton of these events. The lowest occurrence (LO) of Amaurolithus primus is registered at the upper part of the Sanel Mb, below the Tale Gypsum Fm. The LOs of Amaurolithus delicatus and Reticulofenestra rotaria, which mark the base of the Messinian, occur in the lower part of the Chicamo Cycles Fm, above the Tale Gypsum Fm, the Triquetrorhabdulus rugosus-Nicklithus amplificus integrate form and the LO of Nicklithus cf. amplificus in the upper part of the Chicamo Cycles Fm. Taking into account these results, a new calibration of the available magnetostratigraphic data is presented: the Chicamo Cycles Fm were formed during the reverse chron C3Ar and the Tortonian-Messinian boundary should be found within the Tale Gypsum Fm or near the top of the Sanel Mb. The onset of the TSC, the first restriction phase of the Fortuna Basin, is represented by the Fenazar Conglomerate Bed, bottom of the Los Baños Fm, and not by the Tale Gypsum Fm, as previously considered.  相似文献   

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To test vicariant speciation hypotheses derived from geological evidence of the closing of the Tethys Sea, we reconstruct phylogenetic relationships of the predominantly fresh-water killifish genus Aphanius using 3263 aligned base pairs of mitochondrial DNA from samples representing 49 populations of 13 species. We use additional 11 cyprinodontid species as outgroup taxa. Genes analysed include those encoding the partial 12S and 16S ribosomal RNAs; transfer RNAs for valine, leucine, isoleucine, glutamine, methionine, tryptophan, alanine, asparagine, cysteine and tyrosine; and complete nicotinamide adenine dinucleotide dehydrogenase subunit I and II. Molecular substitution rate for this DNA region is estimated at of 8.6 +/- 0.1 x 10(-9) substitutions base pair(-1) year(-1), and is derived from a well dated transgression of the Red Sea into the Wadi Sirhan of Jordan 13 million years ago; an alternate substitution rate of 1.1 +/- 0.2 x 10(-8) substitutions base pair(-1) year(-1) is estimated from fossil evidence. Aphanius forms two major clades which correspond to the former eastern and western Tethys Sea. Within the eastern clade Oligocene divergence into a fresh-water clade inhabiting the Arabian Peninsula and an euhaline clade inhabiting coastal area from Pakistan to Somalia is observed. Within the western Tethys Sea clade we observe a middle Oligocene divergence into Iberian Peninsula and Atlas Mountains, and Turkey and Iran sections. Within Turkey we observe a large amount of genetic differentiation correlated with late Miocene orogenic events. Based on concordance of patterns of phylogenetic relationships and area relationships derived from geological and fossil data, as well as temporal congruence of these patterns, we support a predominantly vicariant-based speciation hypothesis for the genus Aphanius. An exception to this pattern forms the main clade of A. fasciatus, an euhaline circum-Mediterranean species, which shows little genetic differentiation or population structuring, thus providing no support for the hypothesis of vicariant differentiation associated with the Messinian Salinity Crisis. The two phylogenetically deepest events were also likely driven by ecological changes associated with the closing of the Tethys Sea.  相似文献   

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We investigated the phylogeography and speciation of Drilus in the Mediterranean, with focus on the Aegean and especially Crete. Altogether 12 species were sequenced for two fragments of the cytochrome c oxidase subunit I gene (cox1–3′ and cox1–5′ mtDNA) and provided 1381 nucleotides. Both fragments were analysed under the maximum likelihood criterion and Bayesian inference separately and concatenated as a single dataset. The Drilus species from the Peloponnese, the Ionian islands and Crete did not form a monophylum. Drilus sp. E from the Peloponnese and Zakynthos was sister to D. mauritanicus Lucas from Spain in most cases. The remaining Greek Drilus species formed a robustly supported clade in all analyses; however, the species from Crete do not seem to be monophyletic. Estimating species divergences using BEAST, we found out that the key dates in the west Aegean Drilus phylogeography appeared to be the Tortonian Crete–Peloponnese separation (12–9 Mya), the desiccation of the Mediterranean basin during the Messinian Salinity Crisis (5.96–5.33 Mya), and the repeated fragmentation of Crete during the Pliocene and Pleistocene. Within the Drilus lineages, we obtained a substitution rate estimate of 2.75% divergence per million years, which is in excellent agreement with previous studies. A Generalized Mixed Yule Coalescent (GMYC) analysis suggested the presence of six Drilus species in Crete (seven species in total, since the DNA sequences were not available for D. creticus Pic); however, we formally identified and (re)described only four which can be morphologically defined: D. creticus, D. longulus Kiesenwetter, D. horasfakionus sp. nov. and D. baenai sp. nov. These species are endemic to Crete and surrounding islets. Their diagnostic characters are illustrated and an identification key to males of these species is provided. The intraspecific variability, distribution and ecology of all species are discussed and suggestions for further research are given.

http://zoobank.org/urn:lsid:zoobank.org:pub:F8DDBCC1-C2D1-48F7-BE99-55D9ED4C2234  相似文献   

11.
    
An articulated series of centra from latest Miocene deposits of Ukraine is described and identified as a species of Lates. The specimen comes from a locality with an estimated age of 6.04–4.7 Ma, representing the northernmost reach of the Paratethys during the Pontian. The specimen almost certainly represents a distinct species, but because it is incomplete we do not formally name it. The specimen can be differentiated from other species of Lates by features of the centra, including the relative size of the primary lateral fossa of the second centrum, the broadened neural spine of the third centrum, and the relative position of the upper and primary lateral pits on both the fifth and sixth centra. The Ukrainian specimen documents the presence of Lates in the northern Paratethys in the latest Miocene. After this time, species of Lates persisted in freshwaters in the Mediterranean area (Israel and Africa), but are no longer found in the marine waters of the region. We suggest that latines were extirpated from the Mediterranean and Paratethys by the rapidity of the changes in salinity that occurred in the Tethys Mediterranean basin and Paratethys as a result of the Messinian Salinity Crisis.  相似文献   

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The genus Blanus Wagler, 1830 represents limbless, burrowing reptiles of the family Blanidae with disjunct circum-Mediterranean distribution. The recently described species Blanus alexandri Sindaco, Kornilios, Sacchi & Lymberakis, 2014 is known from south-eastern Turkey with a presumed occurrence in the Levant and Iraq. We provide here records from Lebanon and confirm the affiliation of Lebanese populations to this species by mitochondrial and nuclear DNA. Blanus alexandri comprises at least seven deeply evolved phylogenetic clades with up to 11% of uncorrected p-distances in their mitochondrial DNA. This suggests a probably older than the Miocene origin of some of these clades. Populations from Lebanon form a different clade with a genetic diversity that is close to populations from southern Turkey.  相似文献   

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Aim Our aims were: (1) to reconstruct the phylogenetic relationships of daffodils (Narcissus), focusing on the lowland subgenus Hermione and the mountain section Apodanthi; (2) to estimate the temporal setting of diversification; (3) to reconstruct the migration patterns of the lineages; and (4) to examine the microevolutionary differentiation of the wide‐ranging Narcissus tazetta group across the Mediterranean. Location The Mediterranean Basin. Methods Plastid (trnT–L, trnL–F and ndhF) sequences were obtained from 63 populations representing 23 species of Narcissus and combined with published data from 16 species. Phylogenetic relationships and dating were inferred by Bayesian analysis based on geological events and divergence estimates of closely related taxa. A dispersal–extinction–cladogenesis analysis was performed using maximum likelihood methods to infer ancestral geographical distributions, and phylogeographical reconstruction was performed using coalescence analysis. Results Subgenus Hermione is not recognized as a monophyletic group because two of the nine species were found to have a close relationship with the subgenus Narcissus. The results on section Apodanthi confirmed previous findings of its monophyly and phylogenetic relationships within this mountain group. Molecular dating and ancestral range reconstructions suggest that the ancestor of Narcissus originated in the Iberian Peninsula during the Late Oligocene–Early Miocene. Eastward expansion of the lineage range proceeded from the western Mediterranean and involved colonization of mountain ranges in northern Africa. The phylogeography of the N. tazetta group revealed a widespread distribution of certain haplotypes, suggesting wide dispersal and a high level of colonization in the Mediterranean Basin. Main conclusions Our study points to the role of three key historical events in Narcissus diversification: tectonic shifts of the Alboran domain in the western Mediterranean, the Messinian salinity crisis, and the onset of the Mediterranean climate followed by periods of repeated glaciation. Diversification of section Apodanthi probably resulted from allopatric speciation, while subgenus Hermione may have shown more sympatric speciation and high dispersal, despite the lack of apparent adaptations to long‐distance dispersal. This is best exemplified by the presence of both ancestral and recent haplotypes of N. tazetta across the Mediterranean.  相似文献   

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We investigated the phylogeographic patterns of Merodon species (Diptera, Syrphidae) in the Eastern Mediterranean. Ten species were sampled on five different islands and mainland sites as a minimum. All samples were screened for their mtDNA COI barcode haplotype diversity, and for some samples, we additionally generated genomic fingerprints. The recently established zoogeographic distribution categories classify these species as having (1) Balkan distribution; (2) Anatolian distribution; (3) continental areas and large islands distribution; and (4) with wide distribution. The ancestral haplotypes and their geographical localities were estimated with statistical parsimony (TCS). TCS networks identified as the ancestral haplotype samples that originated from localities situated within the distributional category of the species in question. Strong geographical haplotype structuring was detected for many Merodon species. We were particularly interested to test the relative importance of current (Aegean Sea) and past Mid‐Aegean Trench) barriers to dispersal for Merodon flies in the Aegean. We employed phylogenetic β‐diversity (Pβtotal) and its partition in replacement (Pβrepl) and richness difference (Pβrich) to test the importance of each explanatory variable (interisland distance, MAT, and island area) in interisland differences using partial Mantel tests and hierarchical partitioning of variation. β‐Analyses confirmed the importance of both current and past barriers to dispersal on the evolution of group. Current interisland distance was particularly important to explain the replacement of haplotypes, while the MAT was driving differences in richness of haplotypes, revealing the MAT as a strong past barrier whose effects are still visible today in the phylogenetic history of the clade in the Aegean. These results support the hypothesis of a highly restricted dispersal and gene flow among Merodon populations between islands since late Pleistocene. Additionally, patterns of phylogeographic structure deduced from haplotype connections and ISSR genome fingerprinting data revealed a few putative cases of human‐mediated transfers of Merodon spp.  相似文献   

16.
This study proposes to compare the outputs from the CARAIB vegetation model forced by results from the LMD General Circulation Model with interpolated pollen data (Kriging method) from the Mediterranean region during the Messinian. The vegetation maps that have been obtained represent distinct phases of the salinity crisis: before the crisis and during the marginal evaporitic phase (interpolated map), and during the complete desiccation phase (simulated map). However, they are comparable in terms of vegetation density and agree on a strong contrast between the Northern (forest vegetation) and Southern (open vegetation) Mediterranean regions. Main differences concern the type of forests in the northern Mediterranean region, which are explained by discrepancies between precipitation amount predicted by the model and that calculated by a transfer function using pollen records. The interpolation method has been successfully tested in France using interpolated current pollen records by comparison with the present-day potential vegetation map. The resulting Messinian map is useful to validate or improve model simulation which does not take into account the depth of the Mediterranean Basin when it dried up. The Southern Mediterranean landscapes were open, with a steppe-like vegetation to the West and a savannah-like vegetation to the East. Forests prevailed to the North, organized in a mosaic system mainly controlled by relief. Such a contrast provides some explanation of the large number of deep fluvial canyons cut on the Northern margin at opposed to the South during the Mediterranean desiccation.  相似文献   

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AIM: Our aims were to assess the phylogeographic patterns of genetic diversity in eastern Mediterranean water frogs and to estimate divergence times using different geological scenarios. We related divergence times to past geological events and discuss the relevance of our data for the systematics of eastern Mediterranean water frogs. LOCATION: The eastern Mediterranean region. METHODS: Genetic diversity and divergence were calculated using sequences of two protein-coding mitochondrial (mt) genes: ND2 (1038 bp, 119 sequences) and ND3 (340 bp, 612 sequences). Divergence times were estimated in a Bayesian framework under four geological scenarios representing alternative possible geological histories for the eastern Mediterranean. We then compared the different scenarios using Bayes factors and additional geological data. RESULTS: Extensive genetic diversity in mtDNA divides eastern Mediterranean water frogs into six main haplogroups (MHG). Three MHGs were identified on the Anatolian mainland; the most widespread MHG with the highest diversity is distributed from western Anatolia to the northern shore of the Caspian Sea, including the type locality of Pelophylax ridibundus. The other two Anatolian MHGs are restricted to south-eastern Turkey, occupying localities west and east of the Amanos mountain range. One of the remaining three MHGs is restricted to Cyprus; a second to the Levant; the third was found in the distribution area of European lake frogs (P. ridibundus group), including the Balkans. MAIN CONCLUSIONS: Based on geological evidence and estimates of genetic divergence we hypothesize that the water frogs of Cyprus have been isolated from the Anatolian mainland populations since the end of the Messinian salinity crisis (MSC), i.e. since c. 5.5-5.3 Ma, while our divergence time estimates indicate that the isolation of Crete from the mainland populations (Peloponnese, Anatolia) most likely pre-dates the MSC. The observed rates of divergence imply a time window of c. 1.6-1.1 million years for diversification of the largest Anatolian MHG; divergence between the two other Anatolian MHGs may have begun about 3.0 Ma, apparently as a result of uplift of the Amanos Mountains. Our mtDNA data suggest that the Anatolian water frogs and frogs from Cyprus represent several undescribed species.  相似文献   

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The oriental green lizards of the Lacerta trilineata group are widely distributed in Greece, Anatolia, the eastern Mediterranean, the southern Caucasus, and the Zagros mountains in Iran. We studied their phylogeography using three mitochondrial markers with comprehensive sampling from most representatives of the group. Their phylogeny and divergence times (implementing fossil‐based molecular clock calibrations) were inferred using Bayesian methods, and haplotype networks were reconstructed to assess how genetic diversity and current distributional patterns were shaped. According to our phylogenetic analyses, the group constitutes a well‐supported monophylum containing several distinct evolutionary lineages with high haplotype diversity. Vicariance might explain the divergences within most lineages that have accumulated by range restriction and expansion of populations as a result of Quaternary climate oscillations and glacial refugia. However, niche divergence appears to be a major force promoting speciation, and large scale distributional patterns between lineages were shaped earlier by multiple, independent dispersals out of Anatolia during the Pliocene and early Pleistocene. The results of the present study also suggest that the group is in need of a taxonomical revision because the identified lineages and genetic diversity are not congruent with the currently recognized subspecies. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 398–408.  相似文献   

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