The concept of the oviparity-viviparity continuum refers to the wide range in the length of intrauterine egg retention and, hence, in the stage of embryonic development at oviposition existing in squamates. The evolutionary process underlying this continuum may involve not only a lengthening of egg retention in utero, but also a marked reduction in the thickness of the eggshell. The idea that there may exist a negative correlation between the developmental stage reached by the embryo at oviposition and the eggshell thickness within squamates, although supported by the comparison of oviparous vs. viviparous species, has seldom been evaluated by comparing eggshell thickness of oviparous forms with different lengths of intrauterine egg retention. Eggs of two distinct oviparous clades of the lizard Lacerta vivipara were compared. The eggs laid by females from Slovenian and Italian populations have thicker eggshells, contain embryos on average less developed at the time of oviposition, and require a longer incubation period before hatching than the eggs laid by females from French oviparous populations. Our data and several other examples available from the literature support the idea that the lengthening of intrauterine retention of eggs and the shortening of the subsequent external incubation of eggs are associated with reduction in the thickness of the eggshell, at least in some lineages of oviparous squamates. The current hypotheses that may account for this correlation are presented and a few restrictions and refinements to those hypotheses are discussed. In particular, other changes, such as increased vascularization of the oviduct and of the extraembryonic membranes, may play the same role as the decrease of eggshell thickness in facilitating prolonged intrauterine egg retention in squamates. Future studies should also consider the hypothesis that the length of intrauterine retention might directly depend on the extent of maternal-fetal chemical communication through the eggshell barrier.
We describe Vipera walser, a new viper species from the north‐western Italian Alps. Despite an overall morphological resemblance with Vipera berus, the new species is remarkably distinct genetically from both V. berus and other vipers occurring in western Europe and shows closer affinities to species occurring only in the Caucasus. Morphologically, the new species appear to be more similar to V. berus than to its closest relatives occurring in the Caucasus, but can be readily distinguished in most cases by a combination of meristic features as confirmed by discriminant analysis. The extant population shows a very low genetic variability measured with mitochondrial markers, suggesting that the taxon has suffered a serious population reduction/bottleneck in the past. The species is extremely range‐restricted (less than 500 km2) and occurs only in two disjunct sites within the high rainfall valleys of the Alps north of Biella. This new species should be classified as globally ‘endangered’ due to its small and fragmented range, and an inferred population decline. The main near‐future threats to the species are habitat changes associated with reduced grazing, along with persecution and collecting.
Contact zones between two evolutionary lineages are often useful for understanding the process of speciation because the observed genetic pattern reflects the history of differentiation. The Eurasian lacertid lizard Zootoca vivipara is a potentially interesting model for studying the role of reproductive mode in the speciation of squamate reptiles because it has both oviparous (Zootoca vivipara carniolica) and viviparous (Zootoca vivipara vivipara) populations that have recently been shown to be genetically distinct. We studied a newly-discovered syntopic area of these two Zootoca subspecies in the central Italian Alps using genetic markers to investigate the level of introgression between them. Patterns of genetic differentiation in a fragment of the mitochondrial DNA cytb gene and a set of nuclear microsatellites show that the speciation process is complete in this area, with no evidence of recent introgression. Phylogenetic and genotypic divergence suggests that the two subspecies have experienced long independent evolutionary histories, during which genetic and phenotypic differences evolved. The possible roles of biogeography, reproductive mode, and cytogenetic differentiation in this speciation process are discussed.
Genetic variability, one of the main factors that guarantees species persistence, and species’ conservation status are generally evaluated with indices calculated at the present time. Natural history collections might help compare historical and current genetic diversity so to identify major trends. Here we analysed museum specimens of the lizard Zootoca vivipara carniolica, with a specific and stringent protocol for degraded DNA, in order to contrast its past and current genetic variability, using fragments of one mitochondrial DNA gene. Part of the distributional range of Z. v. carniolica (Po Plain, Italy), heavily impacted by human activities, was investigated. We found two previously unknown haplotypes in populations that are extinct today, suggesting the loss of these haplotypes and thus an overall shrinking of genetic variability. We argue that these results, together with the increasing threats posed by climate and land use changes, suggest that specific conservation measures for the persistence of Z. v. carniolica in Northern Italian lowlands have to be considered.
The description of Vipera walser from the Northern Italian Alps as a new species (Ghielmi et al., 2016, Journal of Zoological Systematics and Evolutionary Research, 54, 161) was one of the most unexpected surprises of European herpetology in the 21st century. In mitochondrial (mt) DNA, it is closely related to a group of vipers only present in the Caucasus region and Northeastern Anatolia. However, its morphology is similar to the V. berus populations that inhabit nearby mountains in the Swiss‐Italian Alps, which raises questions on its relationships and status. We thus sequenced five nuclear (nu) genes to determine the position of V. walser relative to V. berus and to the Caucasian/Northeastern Anatolian vipers in nuDNA. We also reanalyzed five previously sequenced mtDNA fragments. NuDNA markers recovered V. walser as closely related to Italian populations of V. berus and not to the Caucasian/Anatolian species, thus contradicting the mtDNA phylogeny. We checked that each of the five mtDNA fragments independently amplified by Ghielmi et al. (2016, Journal of Zoological Systematics and Evolutionary Research, 54, 161) produced individual gene trees compatible with the concatenated mtDNA phylogeny, thus excluding the hypothesis that NUMTs sequencing generated the mtDNA relationships reported by Ghielmi et al. (2016, Journal of Zoological Systematics and Evolutionary Research, 54, 161). Given the low level of nuclear differentiation between V. walser and the Italian population of V. berus, we argue that ancient admixture between V. berus and the ancestral population of V. walser is the most likely explanation for this case of cyto‐nuclear discordance and we discuss the consequences of these results on the systematic status of V. walser.
Improving the knowledge about the distribution of threatened species is a fundamental step forward for implementing effective measures for species conservation. Distribution maps of herpetological species are often inferred from morphological identification but, in some cases, morphology alone is not reliable for assessing the distribution of look-alike species. The common lizard Zootoca vivipara includes several lineages which, to date, cannot be distinguished by any conventional morphological traits. Therefore, a reliable taxonomic identification for the main Zootoca lineages is only feasible with genetic data. In this study, we identify a morphological trait that can be used to distinguish individuals of the parapatric Z. v. vivipara and Z. carniolica, without the support of molecular analyses. This method may provide a cost-effective tool to better depict the distribution of the threatened Z. carniolica and therefore prioritize the conservation of the locations where the oviparous Alpine lineage is found.
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