Seasonal changes in thermal biology of Podarcis lilfordi (Squamata,Lacertidae) consistently depend on habitat traits

During spring and summer, we studied the thermal ecology of two populations of the Balearic lizard, Podarcis lilfordi, from two coastal islets of Menorca (Balearic Islands, Spain): Aire and Colom. We calculated the accuracy of thermoregulation, that is, the extent to which body temperatures are close to species' thermal optima, the thermal quality of the habitat as the proximity of operative temperatures to thermal optima and effectiveness of thermoregulation, as the extent to which accuracy is higher than thermal quality of the habitat. We found that seasonality affects thermoregulation differently, depending on the lizard population. Those effects are consistent for all thermal parameters under study. The effects of seasonality were significantly stronger in Aire than in Colom islet. Many factors may be responsible for this different effect of seasonality, from differences on physiological traits to differences in the environmental conditions of the two islets, as their resource availability, predator pressure or habitat structure. Identifying the factors that boost or inhibit those seasonal changes would be important to understand thermoregulation in lizards. Slight changes on two similar populations can lead to great differences in thermal ecology of conspecific ectotherms.     

Inter-island variation in femoral secretions of the Balearic lizard,Podarcis lilfordi (Lacertidae)

Many lizards use femoral gland secretions in intraspecific communication. Although there is a consistent interspecific variation in chemical composition of secretions, considerable variation is also often found between populations, which may affect conspecifc recognition and lead to speciation processes. Balearic lizards (Podarcis lilfordi) are currently distributed only in several isolated islets offshore of the main islands with different environmental conditions (vegetation, diet, density of population, etc). Also, there is a high genetic variability between populations. We examined whether there was a similar variation in the composition of the femoral secretions of male lizards, and which could be the causes of such variation. By using GC–MS analyses, we found 75 lipophilic compounds in femoral gland secretions of male P. lilfordi from three representative island populations. Main compounds were steroids (94.4%), mainly cholesterol, but we also found alkanes, ketones, waxy esters, squalene, carboxylic acids and their ethyl esters, alcohols and other minor compounds. However, there were clear differences between populations with respect to the number and relative proportions of compounds. Using the patterns of presence and abundance of compounds in secretions it is possible to predict the population of origin of a lizard. We discuss how these differences could be explained considering genetic and environmental differences between populations.     

Cross-species testing of 27 pre-existing microsatellites in Podarcis gaigeae and Podarcis hispanica (Squamata: Lacertidae)

We tested 27 microsatellite loci for cross-species amplification in the lacertids Podarcis gaigeae and Podarcis hispanica. We detected 11 and 15 polymorphic loci in the former and the latter species, respectively. In a larger sample of individuals from a single population of each species, the number of alleles ranged from five to 23 in 10 of the polymorphic loci in P. gaigeae, and between four and 13 in nine of polymorphic loci in P. hispanica. Two locus deviated from Hardy-Weinberg equilibrium in P. hispanica. Between 11 and 16 of the 27 loci also amplified successfully in three other Podarcis species.     

Molecular phylogeny and biogeography of the wall-lizard Podarcis erhardii (Squamata: Lacertidae)

Erhard's wall lizard, Podarcis erhardii (Sauria: Lacertidae), is highly diversified in Greece and especially in the southern Aegean region. Out of the 28 recognized subspecies, 27 are found in Greece from the North Sporades island-complex in the North Aegean (grossly south of the 39th parallel) to the island of Crete in the South. The species exhibits great morphological and ecological plasticity and inhabits many different habitats from rocky islets and sandy shores to mountaintops as high as 2000m. By examining intraspecific variability at a segment of the mitochondrial gene cytochrome b we have found that that extant populations of P. erhardii are paraphyletic. Furthermore, we have found that subspecies previously defined on the basis of morphological characteristics do not correspond to different molecular phylogenetic clades, so that their status should be reconsidered. The DNA based biogeographical and phylogenetic history of Podarcis in Southern Greece is congruent with available paleogeographic data of the region, which supports the view that DNA sequences may be a useful tool for the study of palaeogeography.     

Isolation and characterization of nine microsatellite loci in Podarcis bocagei (Squamata: Lacertidae)

Nine dinucleotide microsatellite loci were developed through an enrichment protocol for Bocage's wall lizard, Podarcis bocagei Seoane 1884, a lacertid endemic to the Iberian Peninsula. Nineteen primer pairs were designed and tested. From these, nine loci yielded satisfactory results and were screened on 15–19 individuals. These loci revealed a high level of polymorphism (8–15 alleles) and heterozygosity (0.611–0.947) and will certainly be useful in the study of population structure and evolutionary history of this species.     

First Record of the Lizard Genus Podarcis Wagler 1830 (Reptilia,Lacertidae) in the Caucasus

Biology Bulletin - Information on the first discovery of the Italian wall lizard, Podarcis siculus campestris (De Betta 1857), in 2019 is provided for Turkan settlement, Baku, Azerbaijan. The...     

Genetic diversity and historical biogeography of the Maltese wall lizard,Podarcis filfolensis (Squamata: Lacertidae)

Podarcis filfolensis is an endemic lizard from the Maltese archipelago. There is evidence of human-mediated decline and even extirpation of some insular populations of this species. However, information about the intraspecific genetic diversity and phylogeographic patterns of this species is limited. Here we analyze genetic markers from a multi-locus dataset (mtDNA, 2,533 bp; nuclear c-mos gene, 353 bp; 11 microsatellites) for individuals from extant populations of P. filfolensis. Despite generally low genetic variability, two main mitochondrial groupings were clearly identified. In general, individuals from the main island of Malta were genetically distinct from those from Gozo, Comino, Cominotto and Small Blue Lagoon Rock, and also from Linosa and Lampione individuals. Three genetic clusters were detected based on microsatellite data: one was found at higher frequency on Malta, while the other two included samples from the remaining islands, showing some concordance with the mtDNA pattern. A time-calibrated Bayesian tree for the principal mitochondrial lineages indicated strong statistical support for two P. filfolensis lineages that originated in the Pleistocene (105.4–869 Ka). We show that these lineages largely meet the criteria for recognition as evolutionary significant units despite some recent admixture (possibly due to recent translocations between islands). Human disturbance, low genetic variability, evidence of bottlenecks and extirpation on one island indicate that a thorough review of the current conservation status of P. filfolensis would be timely.     

Characterization of polymorphic microsatellite markers in the Dalmatian wall lizard Podarcis melisellensis (Squamata: Lacertidae)

We describe polymerase chain reaction primers and amplification conditions for 13 highly polymorphic microsatellite DNA loci isolated from the Dalmatian wall lizard, Podarcis melisellensis. The number of alleles per locus ranged from 12 to 41, with levels of observed heterozygosity between 0.62 and 0.94. Most of these loci were successfully cross-amplified in the closely related species P. sicula, but levels of polymorphism were always lower.     

Diet composition of the lizard P odarcis lilfordi (Lacertidae) on 2 small islands: an individualresource network approach

Despite it is widely accepted that intrapopulation variation is fundamental to ecological and evolutionary processes,this level of information has only recently been included into network analysis of species/population interactions.When done,it has revealed non-random patterns in the distribution of trophic resources.Nestedness in resource use among individuals is the most recurrent observed pattern,often accompanied by an absence of modularity,but no previous studies examine bipartite modularity.We use network analysis to describe the diet composition of the Balearic endemic lizard Podarcis lilfordi in 2 islets at population and individual levels,based on the occurrence of food items in fecal samples.Our objectives are to 1)compare niche structure at both levels,2)characterize niche partition using nestedness and modularity,and 3)assess how size,sex,season,and spatial location influence niche structure.At population-level niche width was wide,but narrow at the level of the individual.Both islet networks were nested,indicating similar ranking of the food preferences among individuals,but also modular,which was partially explained by seasonality.Sex and body size did not notably affect diet composition.Large niche overlap and therefore possibly relaxed competition were observed among females in one of the islets and during spring on both islets.Likewise,higher modularity in autumn suggests that higher competition could lead to specialization in both populations,because resources are usually scarce in this season.The absence of spatial location influence on niche might respond to fine-grained spatio-temporally distribution of food resources.Behavioral traits,not included in this study,could also influence resource partitioning.     

Isolation and characterization of polymorphic microsatellite markers in the wall lizard Podarcis erhardii (Squamata: Lacertidae)

Five polymorphic microsatellite loci (containing di‐ and penta‐nucleotide repeats) were developed for the Erhard's wall lizard, Podarcis erhardii, an endemic species of southern Balkan Peninsula. The number of alleles ranged from five to 17. Observed and expected heterozygosities ranged from 0.28 to 0.92 and 0.57 to 0.91, respectively. These markers will provide a valuable tool for population genetics analyses and can contribute to the evaluation of conservation programs.     

Isolation and characterization of polymorphic microsatellite loci for the Skyros wall lizard Podarcis gaigeae (Squamata: Lacertidae)

Fifteen polymorphic markers were developed from a microsatellite-enriched library for the lizard Podarcis gaigeae. The loci were checked for variability in 68 individuals from a population on the island of Skyros, Greece. The number of alleles ranged from 3 to 23 per locus and expected heterozygosity from 0.29 and 0.94. Most markers were also polymorphic in three closely related Podarcis species, namely P. erhardi, P. taurica and P. milensis. The markers will be used to examine gene flow and differentiation of island and mainland populations of P. gaigeae.     

Molecular survey of morphological subspecies reveals new mitochondrial lineages in Podarcis muralis (Squamata: Lacertidae) from the Tuscan Archipelago (Italy)

Recent analyses of molecular markers have significantly revised the traditional taxonomy of Podarcis species (Squamata: Lacertidae), leading to critically reconsider the taxonomic value of several subspecies described only on morphological bases. In fact, lizards often exhibit high morphological plasticity both at the intra‐specific and the intra‐population level, especially on islands, where phenotypic divergences are mainly due to local adaptation, rather than to evolutionary differentiation. The Common wall lizard Podarcis muralis exhibits high morphological variability in biometry, pholidosis values and colour pattern. Molecular analyses have confirmed the key role played by the Italian Peninsula as a multi‐glacial refuge for P. muralis, pointing out the lack of congruence between mitochondrial lineages and the four peninsular subspecies currently recognized. Here, we analyse a portion of the protein‐encoding cytochrome b gene in the seven subspecies described for the Tuscan Archipelago (Italy), in order to test whether the mitochondrial haplotypes match the morphologically based taxonomy proposed for Common wall lizard. We also compare our haplotypes with all the others from the Italian Peninsula to investigate the presence of unique genetic lineages in insular populations. Our results do not agree completely with the subspecific division based on morphology. In particular, the phylogenetic analyses show that at least four subspecies are characterized by very similar haplotypes and fall into the same monophyletic clade, whereas the other three subspecies are closer to peninsular populations from central Italy. From these results, we conclude that at least some subspecies could be better regarded as simple eco‐phenotypes; in addition, we provide an explanation for the distinctiveness of exclusive lineages found in the archipelago, which constituted a refuge for this species during last glacial periods.     

No trade-off between sprinting and climbing in two populations of the Lizard Podarcis hispanica (Reptilia: Lacertidae).

Biomechanical reflections suggest that lizards that have specialized in running should differ in parts of their morphological design from lizards that have specialized in climbing. Moreover, adaptation in one direction may conflict with performance in the other. We tested this assumed trade-off by measuring climbing and running performance in two subspecies of the Mediterranean lizard Poaarcis hispanica. One subspecies, P. h. atrata , inhabits a number of small islets near the Spanish coast. It is mainly ground dwelling. The other subspecies, P. h. hispanica , was sampled on the mainland of Spain. It frequendy occupies vertical elements within its habitat. Our data do not support the notion of a trade-off between both types of locomotion. Individuals of P. h. hispanica both run and climb faster than those of P. h. atrata. There is no difference between subspecies in ability to cling onto tilted substrates. Predictions on the morphology of both subspecies, inspired by biomechanical arguments, are not supported by our measurements.     

Multilocus genetic diversity and historical biogeography of the endemic wall lizard from Ibiza and Formentera,Podarcis pityusensis (Squamata: Lacertidae)

Two monophyletic sister species of wall lizards inhabit the two main groups of Balearic Islands: Podarcis lilfordi from islets and small islands around Mallorca and Menorca and Podarcis pityusensis from Ibiza, Formentera and associated islets. Genetic diversity within the endangered P. lilfordi has been well characterized, but P. pityusensis has not been studied in depth. Here, 2430 bp of mtDNA and 15 microsatellite loci were analysed from P. pityusensis populations from across its natural range. Two main genetic groupings were identified, although geographical structuring differed slightly between the mtDNA and the nuclear loci. In general, individuals from islets/islands adjacent to the main island of Ibiza were genetically distinct from those from Formentera and the associated Freus islands for both mtDNA and the nuclear loci. However, most individuals from the island of Ibiza were grouped with neighbouring islets/islands for nuclear loci, but with Formentera and Freus islands for the mitochondrial locus. A time‐calibrated Bayesian tree was constructed for the principal mitochondrial lineages within the Balearics, using the multispecies coalescent model, and provided statistical support for divergence of the two main P. pityusensis lineages 0.111–0.295 Ma. This suggests a mid‐late Pleistocene intraspecific divergence, compared with an early Pleistocene divergence in P. lilfordi, and postdates some major increases in sea level between 0.4 and 0.6 Ma, which may have flooded Formentera. The program IMa2 provided a posterior divergence time of 0.089–0.221 Ma, which was similar to the multispecies coalescent tree estimate. More significantly, it indicated low but asymmetric effective gene copy migration rates, with higher migration from Formentera to Ibiza populations. Our findings suggest that much of the present‐day diversity may have originated from a late Pleistocene colonization of one island group from the other, followed by allopatric divergence of these populations. Subsequent gene flow between these insular groups seems likely to be explained by recent human introductions. Two evolutionary significant units can be defined for P. pityusensis but these units would need to exclude the populations that have been the subjects of recent admixture.     

Genetic variation and differentiation in the lizard, Podarcis wagleriana (Reptilia: Lacertidae)

The electrophoretic variation at 26 presumptive gene loci was investigated in populations of Podarcis ivagleriana from Sicily, the Aegadian Islands, and the Aeolian Islands. For interspecific comparison, samples of the closely related lizard P. sicula from the same geographic area were also used. Population heterogeneity analyses carried out by the estimation of F-statistics and Nei's standard genetic distance, showed a high genetic homogeneity within P. sicula, but a noticeable genetic differentiation within P. wagleriana. In the latter species, Nei's D ranged from 0 to 0.212, and this is because the Aeolian populations were quite distinct from those inhabiting Sicily and the Aegadian Islands. Fixed differences identified at three loci (Ck, Ada, Gp-4) contributed to a relatively high value of Nei's standard genetic distance between the two population groups (D= 0.147). This value is very similar to those found comparing pairs of well-recognized biological species included in the genera Podarcis and Lacerta. Estimation of the time of evolutionary divergence shows that the Aeolian and Sicilian populations of P. wagleriana have been isolated geographically for a long time (0.7 Myr according to Nei's formula; 2 Myr according to Sarich's calibration), indicating evolutionary divergence at the species level. Based on genetic and biogeographic data, it is suggested the recognition of full specific status for the Aeolian populations, for which the name P. rajfonei comb. nova (Aeolian wall lizard) is proposed. Electrophoretic data and comparative species distributions suggest that (1) Podarcis sicula recently colonized the Aeolian Islands, and (2) it has competed successfully with P. rajfonei in this area, greatly reducing the range of the latter and causing the extinction of most of its populations. In fact, P. sicula is widespread in the Aeolian Archipelago, while P. rajfonei is confined at present to one large island (Vulcano) and three tiny islands (Strombolicchio, Scoglio Faraglione, La Canna). This can be considered a classic example of competitive exclusion of a native form (P. rajfonei) by a species accidentally introduced by man (P. sicula).     

麻蜥属Eremias的系统学研究进展

由于各种原因, 麻蜥属的分类仍较混乱.本文对麻蜥属的系统学研究历史做了系统回顾, 从属、亚属、种等各级分类水平叙述了其研究历史, 包括属的建立与订正、物种组成、地理分布特征, 总结了通过外部形态和分子等研究手段, 在分类、系统发育、生物地理等研究层次上所取得的成果, 在此基础上提出需要深入研究麻蜥属的系统发育、谱系地理和物种形成机制等, 包括该属分布格局与青藏高原隆升、天山隆升及中亚荒漠化和第四纪冰期的关系.     

Reproductive Biology of the Rock Lizard (Darevskia saxicola,Reptilia, Lacertidae)

Biology Bulletin - Data on the reproduction of the rock lizard (Darevskia saxicola (Eversmann 1834)) in the gorges of Bol’shaya Laba (Karachay-Cherkessia) and Malaya Laba (Krasnodar...