The leukocyte profile has the potential to be a reliable method to measure health conditions and stress in wild animals, but limitations occur because current knowledge on reference intervals is ...largely incomplete, especially because data come from studies on captive animals involving few individuals from single populations. Here we propose a general framework for achieving reliable leukocyte reference intervals, encompassing a set of internal and external factors, potentially affecting the leukogram. To do so, we present a systematic survey of the hematology of the common wall lizard, Podarcis muralis, involving 794 lizards from 54 populations over the whole geographic range of the species in Italy. Reference intervals for white blood cell (WBC) and leukocyte differential count were obtained by using linear mixed models in a Bayesian framework. The application of the procedure clearly showed that both internal (sex and size) and external (latitude and season) factors are a source of variation of leukocyte profile. Furthermore, the leukogram of common wall lizard has a strong variability among populations, which accounts for more than 50% of the whole variation. Consequently, some common assumptions used in studies on captive individuals are no longer supported in wild populations, namely, i) any group of individuals is a representative sample, ii) any population is representative of all others, iii) geographic clines do not occur over the species range, and iv) seasonal variation has limited effects. We encourage researchers aimed at the definition of leukocyte reference intervals for wild populations of reptiles to involve a large number of populations over a wide geographic range in ad hoc statistical models to disentangle local and geographic effects on leukocyte profile variation.
Hybridisation is increasingly recognised as an important cause of diversification and adaptation. Here, we show how divergence in male secondary sexual characters between two lineages of the common ...wall lizard (Podarcis muralis) gives rise to strong asymmetries in male competitive ability and mating success, resulting in asymmetric hybridisation upon secondary contact. Combined with no negative effects of hybridisation on survival or reproductive characters in F1‐hybrids, these results suggest that introgression should be asymmetric, resulting in the displacement of sexual characters of the sub‐dominant lineage. This prediction was confirmed in two types of secondary contact, across a natural contact zone and in two introduced populations. Our study illustrates how divergence in sexually selected traits via male competition can determine the direction and extent of introgression, contributing to geographic patterns of genetic and phenotypic diversity.
Lizards are ideal for studying colour polymorphism, because some species are polymorphic and the morphs often have different ecological or reproductive strategies. We studied the feeding habits of ...six polymorphic populations of Podarcis muralis to test whether morphs differed in their diet. Some taxa were selected in a similar way by all morphs, but selection on other taxa varied and was characteristic of each morph. Diet was most different for the red and yellow morphs. Two hypotheses could explain these differences: active segregation in the trophic niche or active segregation in space dependent on spatial heterogeneity in prey availability. The former is improbable because P. muralis is considered an opportunistic feeder, whereas the latter could occur if the morphs adopted alternative territorial strategies with consequent spatial segregation.
The evolution of sexual dimorphism (SD) is driven by intricate interplays between sexual and natural selection. When it comes to SD variation within populations, however, environmental factors play a ...major role. Sexually selected traits are expected to be strongly dependent on individual body condition, which is influenced by the local environment that individuals experience. As a consequence, the degree of SD may also depend on resource availability. Here, we investigated the potential drivers of SD expression at two sexually dimorphic morphometric traits, body size (snout vent length) and head shape (head geometric morphometrics), in the Italian wall lizard (Podarcis siculus). We assessed the existence of condition- and context-dependent SD across ten islands of the Aeolian archipelago (southern Italy), at within- and among-population scales. We observed strong geographical variation of SD among islands, and tested three potential SD predictors related to resource availability (individual body condition, ecosystem productivity, temperature). Body condition and ecosystem productivity were the main drivers of body size SD variation, and body condition was also the main driver for head shape SD. Our results highlight that the expression of SD in the Italian wall lizard is both condition- and context-dependent. These results are congruent at within- and among-populations scales highlighting that spatial multi-scale analysis represents a useful approach to understand patterns of SD expression.
Abstract
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.
Abstract
The Mediterranean Basin has experienced extensive change in geology and climate over the past six million years. Yet, the relative importance of key geological events for the distribution ...and genetic structure of the Mediterranean fauna remains poorly understood. Here, we use population genomic and phylogenomic analyses to establish the evolutionary history and genetic structure of common wall lizards (Podarcis muralis). This species is particularly informative because, in contrast to other Mediterranean lizards, it is widespread across the Iberian, Italian, and Balkan Peninsulas, and in extra-Mediterranean regions. We found strong support for six major lineages within P. muralis, which were largely discordant with the phylogenetic relationship of mitochondrial DNA. The most recent common ancestor of extant P. muralis was likely distributed in the Italian Peninsula, and experienced an “Out-of-Italy” expansion following the Messinian salinity crisis (∼5 Mya), resulting in the differentiation into the extant lineages on the Iberian, Italian, and Balkan Peninsulas. Introgression analysis revealed that both inter- and intraspecific gene flows have been pervasive throughout the evolutionary history of P. muralis. For example, the Southern Italy lineage has a hybrid origin, formed through admixture between the Central Italy lineage and an ancient lineage that was the sister to all other P. muralis. More recent genetic differentiation is associated with the onset of the Quaternary glaciations, which influenced population dynamics and genetic diversity of contemporary lineages. These results demonstrate the pervasive role of Mediterranean geology and climate for the evolutionary history and population genetic structure of extant species.
Colour variability is largely widespread in the animal world as it is tightly associated with fitness and survivorship. Therefore, the drivers and implications of such variability have been of great ...interest for zoologists in the past decades. Reptiles are excellent models to investigate colour variations and expression under different conditions. Here, we focused on melanism occurrence in the two main lineages of
Hierophis viridiflavus
at the scale of the species distribution, by extracting available data from iNaturalist, a citizen science network, with the aim of detecting any pure effect of climate or local habitat on colour expression. Our analyses highlighted that habitat does not explain differences in phenotypes, whereas marked effects of geographic and climatic variables were detected. However, the observed climatic effects could be a proxy of the geographical distribution of the two groups, and thus the high occurrence of bright colourations in western populations of the eastern lineage could be addressed to an ongoing event of asymmetric gene flow in contact zones. The current distribution of phenotypes could be the outcome of the evolutionary history of the species combined with the geological history of the Mediterranean region. This investigation, though, is only preliminary and molecular analyses on highly variable regions of the genome are mandatory to address this issue.
Ecological aspects of syntopic geckoes were rarely addressed in the Mediterranean basin. We reported basic information on habitat use, and activity patterns of three species found in syntopy in Cala ...Violina site (divided in three subareas), a highly touristic beach located in southern Tuscany, central Italy, during 2009-2010. The most abundant species at first capture is Hemidactylus turcicus (94 individuals), while Tarentola mauritanica and Euleptes europaea are less represented (28 animals in both cases). Total captures and recaptures were 175. Sex ratio did not differ from 1:1 in all the species, nor sexes of adults did differ in size. Ambient temperatures did not differ in T. mauritanica and E. europaea, while were different in H. turcicus. Despite the humidity of capture sites did not vary among species, we recorded the highest number of E. europaea at 95% and H. turcicus at 62% humidity. Wind influenced negatively T. mauritanica and H. turcicus presence, not on E. europaea. Higher observation rate took place between 21:00 and 22:00. After 23:00, only Euleptes was active. Height from the ground was different only in H. turcicus. General Linear Models showed that interaction substrate-height at capture was important for Euleptes, not for the other two species. Along the area, E. europaea was more concentrated in the northern patch, while T. mauritanica and H. turcicus distributed more homogeneously. We suggest limitation of human presence for conservation purposes.
Abstract
Animal communication depends on signals conveying information to a receiver who must perceive and decode them. Signals involved in territoriality are usually complex stimuli that should be ...correctly interpreted to avoid unnecessary conflicts. Lacertids use both visual and chemical stimuli in modulating their aggressive response against conspecifics and the rival’s size is one of the most important information, affecting the success probability in combat. To assess the actual ability of decoding information about a rival’s size based on its chemical stimulus alone, 60 males of Podarcis muralis were tested for three consecutive days in an arena bearing a mirror (to simulate an equal-sized intruder), and the chemical cues (femoral secretions) from an unknown individual of different size. Significant differences were observed in tongue-flicks number, which grew as the size difference between the focal lizard and the secretion donor decreased. This can be interpreted as the need for the lizard to better evaluate the potential competitor’s characteristics. The size difference also affected the number of bites against the mirror. They increased when the size of the focal lizard was larger than the donor triggering the aggressive response with a higher probability of winning the contest. This confirms that the focal lizard had correctly decoded the information about the opponent’s size by chemical stimulus. Although previous studies have shown that some components of the chemical signals are potentially informative about the signaler’s size, this is the first demonstration that male P. muralis is actually able to decode and use such information.
Lizards communicate through chemical signals, which convey information about the signaller’ quality and identity. We demonstrate that males can perceive and decode information about actual rival’s size using chemical cues, even when they see a size-matched mirrored image. Individuals attack smaller rivals more often, but spend the most time assessing rivals when their opponents are of similar size, and less time assessing rivals when they are substantially smaller or larger than them. This could be useful to avoid unnecessary and costly fightings with stronger opponents.
Animals modulate intraspecific signal shape and intensity, notably during reproductive periods. Signal variability typically follows a seasonal scheme, traceable through the expression of visual, ...acoustic, chemical and behavioral patterns. The chemical channel is particularly important in lizards, as demonstrated by well-developed epidermal glands in the cloacal region that secrete lipids and proteins recognized by conspecifics. In males, the seasonal pattern of gland activity is underpinned by variation of circulating androgens. Changes in the composition of lipid secretions convey information about the signaler’s quality (e.g., size, immunity). Presumably, individual identity is associated with a protein signature present in the femoral secretions, but this has been poorly investigated. For the first time, we assessed the seasonal variability of the protein signal in relation to plasma testosterone level (T), glandular activity and the concentration of provitamin D
3
in the lipid fraction. We sampled 174 male common wall lizards (
Podarcis muralis
) over the entire activity season. An elevation of T was observed one to two months before the secretion peak of lipids during the mating season; such expected delay between hormonal fluctuation and maximal physiological response fits well with the assumption that provitamin D
3
indicates individual quality. One-dimensional electrophoretic analysis of proteins showed that gel bands were preserved over the season with an invariant region; a result in agreement with the hypothesis that proteins are stable identity signals. However, the relative intensity of bands varied markedly, synchronously with that of lipid secretion pattern. These variations of protein secretion suggest additional roles of proteins, an issue that requires further studies.
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