The evolution of viviparity in squamates has been the focus of much scientific attention in previous years. In particular, the possibility of the transition from viviparity back to oviparity has been the subject of a vigorous debate. Some studies have suggested this reversal is more frequent than previously thought. However, none of them provide conclusive evidence. We investigated this problem by studying the phylogenetic relationships between oviparous and viviparous lineages of the reproductively bimodal lizard species Zootoca vivipara . Our results show that viviparous populations are not monophyletic, and that several evolutionary transitions in parity mode have occurred. The most parsimonious scenario involves a single origin of viviparity followed by a reversal back to oviparity. This is the first study with a strongly supported phylogenetic framework supporting a transition from viviparity to oviparity.
The European common lizard, Zootoca vivipara, is the most widespread terrestrial reptile in the world. It occupies almost the entire Northern Eurasia and includes four viviparous and two oviparous lineages. We analysed how female snout-vent length (SVL), clutch size (CS), hatchling mass (HM), and relative clutch mass (RCM) is associated with the reproductive mode and climate throughout the species range and across the evolutionary lineages within Z. vivipara. The studied variables were scored for 1,280 females and over 3,000 hatchlings from 44 geographically distinct study samples. Across the species range, SVL of reproductive females tends to decrease in less continental climates, whereas CS corrected for female SVL and RCM tend to decrease in climates with cool summer. Both relationships are likely to indicate direct phenotypic responses to climate. For viviparous lineages, the pattern of co-variation between female SVL, CS and HM among populations is similar to that between individual females within populations. Consistent with the hypothesis that female reproductive output is constrained by her body volume, the oviparous clade with shortest retention of eggs in utero showed highest HM, the oviparous clade with longer egg retention showed lower HM, and clades with the longest egg retention (viviparous forms) had lowest HM. Viviparous populations exhibited distinctly lower HM than the other European lacertids of similar female SVL, many of them also displaying unusually high RCM. This pattern is consistent with Winkler and Wallin’s model predicting a negative evolutionary link between the total reproductive investment and allocation to individual offspring.
We provide an integrative analysis of the diversity of the E. multiocellata—E. przewalskii species complex in Central and Middle Asia using morphological and molecular (COI DNA-barcoding) data. We report preliminary data on mtDNA variation within this group and clarify the taxonomic status and distribution of the members of the species complex. We also provide a description of a new Eremias species from Eastern Kazakhstan and western Mongolia, where it occurs in sympatry with E. multiocellata sensu stricto, from which it can be clearly differentiated using both morphological and molecular characters. The new species, described as Eremias dzungarica sp. nov., is assigned to the subgenus Pareremias on the basis of the following features: subocular not reaching mouth edge; one frontonasal; two supraoculars; the row of small granular scales between supraoculars and frontal with frontoparietals absent; distance between the femoral pore rows being wide; femoral pore rows not reaching knee-joint; coloration pattern with light colored ocelli with black edging. The new species can be distinguished from its congeners on the basis of the following morphological attributes: a medium-sized lacertid lizard, maximum snout-vent length (SVL) = 64.5 mm, tail being ca. 1.5 times longer than body length (SVL), hindlimbs relatively long (hindlimb length to SVL ratio 0.46); subocular scale not reaching mouth edge, in touch with 6–8 supralabials; males with bright coloration consisting of 2–3 dorsolateral rows of light-colored ocelli with thick black edging; the ventral row of ocelli in life is greenish to bluish; dorsal pattern consisting of black irregular blotches along the middorsal line. We also report on the high genetic and morphological diversity of E. multiocellata in Mongolia and China, synonymize E. m. bannikowi with the nominative form E. m. multiocellata, discuss variation within E. przewalskii, synonymize E. p. tuvensis with the nominative form E. przewalskii, provide new data on E. cf. reticulata and E. m. tsaganbogdensis, confirm validity and clarify distribution ranges of E. stummeri, E. szczerbaki and E. yarkandensis and discuss further progress on taxonomic studies of the E. multiocellata—E. przewalskii species complex.
Abstract. The snout-vent length (SVL), a conventional measure of overall body size in lizards and snakes, is used in a wide variety of ecological, evolutionary, and taxonomical studies. Trends in SVL variation are often analysed using data from several researchers (observers), but possible confounding effects due to inter-observer differences in measurement protocols have never been appropriately examined. This study reports inter-observer biases between eleven herpetologists who measured the same specimens of the Eurasian common lizards (21 adult specimens were examined by eight observers and additional 192 specimens by two observers). Intra-observer bias over time (1.5-15 months between measuring sessions) was also estimated. In the vast majority of comparisons, mean difference between the first author and another observer varied from −1.0 to +0.8 mm, or from −1.9 to +1.6% if expressed as a percent of the specimen's SVL value. Some non-regular effects of sex and study sample on the studied bias were revealed, and their possible reasons are discussed. We are advising the researchers who intensively collect SVL and other morphometric data to consider testing intra-observer and inter-observer biases and to establish etalon samples available for re-examinations.
Reproductive mode, ancestry, and climate are hypothesized to determine body size variation in reptiles but their effects have rarely been estimated simultaneously, especially at the intraspecific level. The common lizard (Zootoca vivipara) occupies almost the entire Northern Eurasia and includes viviparous and oviparous lineages, thus representing an excellent model for such studies. Using body length data for >10,000 individuals from 72 geographically distinct populations over the species' range, we analyzed how sex‐specific adult body size and sexual size dimorphism (SSD) is associated with reproductive mode, lineage identity, and several climatic variables. Variation in male size was low and poorly explained by our predictors. In contrast, female size and SSD varied considerably, demonstrating significant effects of reproductive mode and particularly seasonality. Populations of the western oviparous lineage (northern Spain, south‐western France) exhibited a smaller female size and less female‐biased SSD than those of the western viviparous (France to Eastern Europe) and the eastern viviparous (Eastern Europe to Far East) lineages; this pattern persisted even after controlling for climatic effects. The phenotypic response to seasonality was complex: across the lineages, as well as within the eastern viviparous lineage, female size and SSD increase with increasing seasonality, whereas the western viviparous lineage followed the opposing trends. Altogether, viviparous populations seem to follow a saw‐tooth geographic cline, which might reflect the nonmonotonic relationship of body size at maturity in females with the length of activity season. This relationship is predicted to arise in perennial ectotherms as a response to environmental constraints caused by seasonality of growth and reproduction. The SSD allometry followed the converse of Rensch's rule, a rare pattern for amniotes. Our results provide the first evidence of opposing body size—climate relationships in intraspecific units.
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