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.
T. U. (2007). Phylogeography and cryptic variation within the Lacerta viridis complex (Lacertidae, Reptilia). -Zoologica Scripta, 36 , 119-131. It is well known that the current genetic pattern of many European species has been highly influenced by climatic changes during the Pleistocene. While there are many well known vertebrate examples, knowledge about squamate reptiles is sparse. To obtain more data, a range-wide sampling of Lacerta viridis was conducted and phylogenetic relations within the L. viridis complex were analysed using an mtDNA fragment encompassing part of cytochrome b , the adjacent tRNA genes and the noncoding control region. Most genetic divergence was found in the south of the distribution range. The Carpathian Basin and the regions north of the Carpathians and Alps are inhabited by the same mitochondrial lineage, corresponding to Lacerta viridis viridis . Three distinct lineages occurred in the south-eastern Balkanscorresponding to L. v. viridis , L. v. meridionalis , L. v. guentherpetersi -as well as a fourth lineage for which no subspecies name is available. This distribution pattern suggests a rapid range expansion of L. v. viridis after the Holocene warming, leading to a colonization of the northern part of the species range. An unexpected finding was that a highly distinct genetic lineage occurs along the western Balkan coast. Phylogenetic analyses (Bayesian, maximum likelihood, maximum parsimony) suggested that this west Balkan lineage could represent the sister taxon of Lacerta bilineata . Due to the morphological similarity of taxa within the L. viridis complex this cryptic taxon was previously assigned to L. v. viridis . The distribution pattern of several parapatric, in part highly, distinct genetic lineages suggested the existence of several refuges in close proximity on the southern Balkans. Within L. bilineata sensu stricto a generally similar pattern emerged, with a high genetic diversity on the Apennine peninsula, arguing for two distinct refuges there, and a low genetic diversity in the northern part of the range. Close to the south-eastern Alps, three distinct lineages ( L. b. bilineata , L. v. viridis , west Balkan taxon) occurred within close proximity. We suggest that the west Balkan lineage represents an early offshoot of L. bilineata that was isolated during a previous Pleistocene glacial from the more western L. bilineata populations, which survived in refuges on the Apennine peninsula.
The fecundity-advantage hypothesis (FAH) explains larger female size relative to male size as a correlated response to fecundity selection. We explored FAH by investigating geographic variation in female reproductive output and its relation to sexual size dimorphism (SSD) in Lacerta agilis, an oviparous lizard occupying a major part of temperate Eurasia. We analysed how sex-specific body size and SSD are associated with two putative indicators of fecundity selection intensity (clutch size and the slope of the clutch size-female size relationship) and with two climatic variables throughout the species range and across two widespread evolutionary lineages. Variation within the lineages provides no support for FAH. In contrast, the divergence between the lineages is in line with FAH: the lineage with consistently female-biased SSD (L. a. agilis) exhibits higher clutch size and steeper fecundity slope than the lineage with an inconsistent and variable SSD (L. a. exigua). L. a. agilis shows lower offspring size (egg mass, hatchling mass) and higher clutch mass relative to female mass than L. a. exigua, that is both possible ways to enhance offspring number are exerted. As the SSD difference is due to male size (smaller males in L. a. agilis), fecundity selection favouring larger females, together with viability selection for smaller size in both sexes, would explain the female-biased SSD and reproductive characteristics of L. a. agilis. The pattern of intraspecific life-history divergence in L. agilis is strikingly similar to that between oviparous and viviparous populations of a related species Zootoca vivipara. Evolutionary implications of this parallelism are discussed.
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