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Die landgebundenen Wirbeltiere des festländischen Iberiens (Süßwasserfische, Amphibien, Reptilien mit Ausnahme der Meeresschildkröten, Säugetiere mit Ausnahme der Chiroptera, Pinnipedia und Cetacea) bilden, wie im einzelnen gezeigt wird, in ihrer Gesamtheit eine besondere faunistische Einheit: ungefähr die Hälfte der Formen (Spezies oder Subspezies) ist endemisch; das gilt für 73 % der echten Süßwasserfische, 47 % der Amphibien, 36 % der Reptilien und 53 % der Säugetiere. Die geographische Isolierung durch die Pyrenäen und das Mittelmeer ist gewiß die Hauptursache für die Entwicklung einer so eigenartigen Fauna, aber auch andere Faktoren (die sehr verschiedenen Klimate und Biotope innerhalb Iberiens, der im Vergleich mit anderen Gebieten Europas geringere Einfluß der Vereisungen usw. haben sicher ebenfalls eine sehr wichtige Rolle gespielt. Acanthodactylus erythrurus erythrurus, Algyroides hidalgoi, Algyroides marchi marchi, Algyroides marchi niethammeri, Lacerta agilis agilis, Lacerta hispanica hispanica, Lacerta hispanica bocagei, Lacerta hispanica vaucheri, Lacerta lepida lepida, Lacerta lepida nevadensis, Lacerta monticola monticola, Lacerta monticola cantabrica, Lacerta monticola cyreni, Lacerta schreiberi, Lacerta viridis viridis, Lacerta vivipara, Psammodromus algirus algirus, Psammodromus hispanicus hispanicus, Psammodromus hispanicus edwardsianus.

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The determination of amphibians and reptiles species, their altitudinal range and biotopes in Les Planes de Son and the Mata de València, was determined by sampling the study area from autumn 2006 to spring of 2007. Compared to other studied Pyrenean areas, the herpetofauna is relatively poor. We only found 5 species of amphibians and 9 of reptiles. Specific samplings discard the presence of two Pyrenean endemics, de newt (Calotriton asper) and the lizard (Iberolacerta bonnali). Interestingly, we found an altitudinal replacement of ocelated lizard (Timon lepidus) by green lizard (Lacerta bilineata) and of Iberian rocky lizard (Podarcis hispanica) by Podarcis muralis. Less forested habitats situated at low altitude are the richest in amphibian and reptile species, in contrast to the highest places inhabited by few species (Rana temporaria, Podarcis muralis and Vipera aspis).

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Es gibt sicherlich verschiedene Möglichkeiten, im Terrarium gehaltene Eidechsen zu überwintern. Der eine schwört auf seinen Keller, der andere auf eine laubgefüllte Grube in seinem Garten. Nun, wem beides nicht zur Verfügung steht, dem kann auch ein Kühlschrank ganz ausgezeichnete Dienste leisten. Freilich ist nicht gemeint, die Eidechsen frei zwischen Wurst und Käse zu betten, vielmehr bedarf es einiger Vorbereitungen. Im wesentlichen stellen sich zur Überwinterung von Eidechsen 4 Fragen: 1. wen? 2. wann? 3. wie? Und 4. wie lange? Auf diese 4 Fragen soll im in diesem Bericht eine Antwort gegeben werden.

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The Columbretes archipelago consists of a group of small volcanic islets located in the western Mediterranean near the east of the Iberian Peninsula. Four of its islands are inhabited by the wall lizard Podarcis liolepis, whose populations have been considered vulnerable. The purpose of this study was to assess the level of genetic diversity across the archipelago and the possible evolutionary origin of the Columbretes lizard populations. Additionally, we investigated the evolu-tionary ecology of these populations using a DNA-based metabarcoding approach to characterise both their microbiota and trophic interactions. The genetic results reported very low genetic diversity and corroborated the conspecificity between insular populations and P. liolepis from the mainland (Peñagolosa region). The results of the metabarcoding analyses based on faecal samples were in accordance with an omnivorous ecology, suggesting that specific microbiota communities in the insular populations might be correlated with differences in host ecology and phylogeny. These results are a valuable contribution to the current understanding of the evolution of Columbretes’ lizards and provide important information for conservation management.

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The phylogenetic relationships among the wall lizards of the Podarcis hispanicus complex that inhabit the south-east (SE) of the Iberian Peninsula and other lineages of the complex remain unclear. In this study, four mitochondrial and two nuclear mark-ers were used to study genetic relationships within this complex. The phylogenetic analyses based on mtDNA gene trees constructed with ML and BI, and a species tree using *BEAST support three divergent clades in this region: the Valencia, Galera and Albacete/Murcia lineages. These three lineages were also corroborated in spe-cies delimitation analyses based on mtDNA using bPTP, mPTP, GMYC, ABGD and BAPS. Bayesian inference species delimitation method (BPP) based on both nuclear data and a combined data set (mtDNA + nuclear) showed high posterior probabilities for these three SE lineages (≥0.94) and another Bayesian analysis (STACEY) based on combined data set recovered the same three groups in this region. Divergence time dating of the species tree provided an estimated divergence of the Galera line-age from the other SE group (Podarcis vaucheri, (Albacete/Murcia, Valencia)) at 12.48 Ma. During this period, the Betic–Rifian arc was isolated, which could have caused the isolation of the Galera form distributed to the south of the Betic Corridor. Although lizards from the Albacete/Murcia and Galera lineage are morphologically similar, they clearly represent distinct genetic lineages. The noteworthy separation of the Galera lineage enables us to conclude that this lineage must be considered as a new full species.

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A complete catalogue is provided for the type specimens of lacertid lizards in the herpetological collection of the Zoological Museum, Berlin, as of December, 1993. The collection contains a total of 514 type specimens, representing 63 taxa, of which 42 are currently regarded as valid at the specific or subspecific level. Types representing an additional four taxa appear to have been lost from the collection. The collections are especially rich in African lacertid types and in historically significant specimens collected early in the Nineteenth Century by Pallas, Eversmann, and Hemprich & Ehrenberg. Also present are the type series of many subspecies of Podarcis lilfordi and P. pityusensis described by Eisentraut.

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The life history of an organism is the combination of age-specific survival probabilities and fecundities it displays in its natural environment. Hence, an organism`s life history is characterised by its age and size at maturity, frequency of reproduction, clutch or litter size, size of eggs and hatchlings, and survivorship at different life stages (as embryos, neonates, immatures, adults). Variation in these traits can be studied at different levels: within a population, among populations of a single species, and among different species. I here give an overview of studies that explored life history variation in European lacertid lizards.

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Oganismal performance abilities occupy a central position in phenotypic evolution; they are determined by suites of interacting lower-level traits (e.g., morphology and physiology) and they are a primary focus of natural selection. The mechanisms by which higher levels of organismal performance are achieved during evolution are therefore fundamentally important for understanding correlated evolution in general and coadaptation in particular. Here we address correlated evolution of morphological, physiological, and behavioral characteristics that influence interspecific variation in sprint speed in a clade of lacertid lizards. Phylogenetic analyses using independent contrasts indicate that the evolution of high maximum sprinting abilities (measured on a photocell-timed racetrack) has occurred via the evolution of (1) longer hind limbs relative to body size, and (2) a higher physiologically optimum temperature for sprinting. For ectotherms, which experience variable body temperatures while active, sprinting abilities in nature depend on both maximum capacities and relative performance levels (i.e., percent of maximum) that can be attained. With respect to temperature effects, relative performance levels are determined by the interaction between thermal physiology and thermoregulatory behavior. Among the 13 species or subspecies of lizards in the present study, differences in the optimal temperature for sprinting (body temperature at which lizards run fastest) closely matched interspecific variation in median preferred body temperature (measured in a laboratory photothermal gradient), in- dicating correlated evolution of thermal physiology and thermal preferences. variability of the preferred body tem- peratures maintained by each species is, across species, negatively correlated with the thermal-performance breadth (range of body temperatures over which lizards can run relatively fast). This pattern leads to interspecific differences in the levels of relative sprint speed that lizards are predicted to attain while active at their preferred temperatures. The highest levels of predicted relative performance are achieved by species that combine a narrow, precise distribution of preferred temperatures with the ability to sprint at near-maximum speeds over a wide range of body temperatures. The observed among-species differences in predicted relative speed were positively correlated with the interspecific variation in maximum sprinting capacities. Thus, species that attain the highest maximum speeds are (1) also able to run at near-maximum levels over a wide range of temperatures and (2) also maintain body temperatures within a narrow zone near the optimal temperature for sprinting. The observed pattern of correlated evolution therefore has involved traits at distinct levels of biological organization, that is, morphology, physiology, and behavior; and trade- offs are not evident. We hypothesize that this particular trait combination has evolved in response to coadaptational selection pressures. We also discuss our results in the context of possible evolutionary responses to global climatic change.

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The list of amphibians and reptiles living in the Provence-Alpes-Côte d’Azur constitutes an inventory of up-to-date knowledge on herpetofauna. It contains 22 amphibian taxons and 41 reptile taxons. It also contains their regional distribution, a list of the relevant national, European, and international protection laws, as well as the conservation status of each species.

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The list of amphibians and reptiles living in the Provence-Alpes-Côte d’Azur constitutes an inventory of up-to-date knowledge on herpetofauna.

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Short note on a concolor morph of Podarcis liolepis liolepis (BOULENGER, 1905) in the old botanical garden (Montjuïc) of Barcelona.

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Geographic range of Podarcis liolepis sebastiani (Klemmer, 1964) (Squamata, Lac­ertidae) in France. The Catalonian wall lizard, Podarcis liolepis, is a Mediterranean species that can be mainly found in France around the Mediterranean belt. In Aquitaine, the subspecies Podarcis liolepis sebastiani enters from Spain on the French Basque country slope. Until now, studies on French ter- ritory have restricted the species to a few localities along the Spanish border. Since spring 2010, specif- ic surveys have been undertaken in the scope of the Atlas of amphibians and reptiles of Aquitaine. Based on our results, the species is widespread in the French Basque country as far as 30 km north from the Spanish-French border. Moreover, the presence of the Catalonian wall lizard has been confirmed in the East of the department of Pyrénées-Atlantiques, in the Aspe Valley. This latter finding suggests new interesting hypotheses about the ancient colonization paths of the species on the northern slope of the Western Pyrenees.

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In 2015 the author found several lizards of this complex, of which he was not able to determine the correct spercies name. The males had no pattern (concolor), the females had the normal dorsolateral stripes for this group. The animals were quite small and slender. Consultation with some herpetologists did not provide more clarity. Five years later the author received an e-mail from one of them (GENIEZ, pers. comm.), that it was probably a Catalonian Wall lizard, Podarcis liolepis. Concolor morphs have been described for this species, but these animals are larger in size and occur much further north. However, genomic data have confirmed that there is a southern lineage, clearly P. liolepis, but distinct from the northern animals, that is smaller in size and more flattened.

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Hardenberg Castle north of Göttingen, Germany harbours two introduced species of wall lizard, the Eurpean Wall Lizard Podarcis muralis brogniardii and the Catalonian Wall Lizards Podarcis liolepis. The genetic study by Schulte et al. (2012a) has found that the European Wall Lizards belong the the Western France clade and that the Catalonian Wall Lizards come from near Girona in NE Spain, thus representing the nominal subspecies, and that the import was from the Eastern central Pyrenees. A visit to the locality in October 2019 throws some doubt on this, as both species seem to be mixed up from various sources, the new populations being harder to distinguish fom each other than their natural ancestors in case of syntopy. It is suggested that there have been two sources for each species, namely that the European Wall Lizards are a mixture with animals of the Eastern France clade, and the Catalonian Wall Lizards have been mixed with specimens from the Northwestern Pyrenees belonging to the subspecies sebastiani. Observations on behaviour, habitat preference, and relative abundance of the species are also reported, and the criteria for distinction of the two species at Hardenberg castle are tested.

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This thesis comprises part of a wider project that addresses the evolution of Mediterranean lizards of the genus Podarcis (Wagler, 1830), and is mainly focused on insular populations. This genus is widespread across the Mediterranean basin. They exhibit a great intraspecific variability in morphological traits. In particular, melanic and non-melanic populations (which are completely green or brown, and/or with intermediate coloration) are found on different islands. The causes for this variability remain unknown. The Podarcis genus contains various insular endemism such as Podarcis tiliguerta from Corsica and Sardinia islands, and P. lilfordi and P. pityusensis from the Balearic archipelago. The Mediterranean basin is considered a biodiversity hotspot due to the presence of an exceptional number of endemic species and Podarcis provides an important component of this biodiversity. It can provide information on the effects of fragmentation and/or degradation of habitat. Analyses of Podarcis also have important implications for conservation of the group itself: understanding the evolutionary framework provides a basis for future conservation actions. The present thesis is focused on the molecular study of the evolution of several Mediterranean Podarcis. It aims to build on and extend previous studies that made evolutionary inferences from just the mitochondrial DNA. This is achieved by analysing several nuclear markers (MC1R, RAG1, APOBE28, BLC9L, KIAA2018 and KIF24) in combination with mtDNA data. The main aim is to address two important evolutionary questions: i) Factors determining melanism in insular populations P. lilfordi and P. pityusensis present an important colour variation and polymorphism in the MC1R gene has been related to pigmentation in other species. Therefore, the relationship between DNA substitutions and melanism in different populations was studied. We did not detect any differences in the MC1R sequence that we analysed and so this sequence does not appear to be the basis of the colour variation in these Balearic Podarcis. Differences in gene expression were also compared between melanic and nonmelanic individuals, but no differences that were directly related to MC1R expression were detected. Nevertheless, the transcriptome profiles of melanic and non-melanic individuals from two populations of P. lilfordi did show some differences. The main differences found between the two groups were: non-melanic samples showed overexpression of stress-response genes, while melanic individuals had higher metabolic rates and an increased inflammatory response to exogenous antigens. Despite finding differences in expression, our study was unable to reveal major insights into the genetic basis of melanism in these island lizards. ii) Phylogeny and phylogeography of different endemic species. This work contributes important findings relevant to the systematics and evolution of the species P. tiliguerta endemic to Corsica and Sardina. Using Bayesian phylogenetic dating, we determined that this species originated ~11 Ma ago. Both markers (nuclear genes and mtDNA) showed a high level of intraspecific divergence, especially between Corsica and Sardinia. Ancient divergence (which was estimated at 9.5 Ma ago) and associated high levels of between-island lineage sorting in nuclear markers support the view that Corsican and Sardinian populations should be recognized as two different species. Podarcis were also studied from the Columbretes archipelago, a small island group off the coast east of Spain. Populations were previously recognized as a subspecies of P. hispanica or as P. atrata species. However, the suitability of this taxonomic designation has been made unclear by recent systematics studies of Podarcis that have led to major taxonomic changes. We used nuclear and mitochondrial genes to define their phylogenetic relationships. Columbretes populations were found to be most closely related to the new taxon P. liolepis.

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Mitochondrial DNA analysis indicates that Podarcis vaucheri is a species complex with one clade inhabiting both north and south shores of the Strait of Gibraltar and one clade restricted to North Africa. While each clade exhibits differing morphology, allozyme analysis suggests varying degrees of reproductive contact among populations within clades. Ancestral P. vaucheri appears to have been restricted to insular areas of the Betic-Rif Massif and mainland Africa during Miocene, with major lineage differentiation occurring during Pliocene. Our analysis further confirms that Podarcis atrata warrants species status and suggests that it may be more widely distributed than previously reported.

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The process of speciation is often equated to the acquisition of reproductive isolation. This is a process typically envisioned as being correlated to divergence time: as time progresses the more likely it will be for diverging taxa to experience restrictions to gene exchange. This correlation has been tested and verified in a wide range of taxa; however, other aspects of differentiation, such as ecological divergence, have also been shown to correlate with reproductive isolation independently of divergence time or genomic differentiation. Characterizing reproductive isolation in multiple pairs of closely related taxa that are in different stages of the divergence process is a way of addressing these questions and disentangling the factors are more importante as engines of reproductive isolation. Wall lizards of genus Podarcis, particularly those belonging to the Iberian and North African group, are an appropriate model for this type of evaluations since they comprise various differentiated species with that establish a variety of natural contact zones. In this study we sampled and characterized patterns of hybridization in 9 contact zones between different species pairs using RADseq data. We found a generally low proportion of hybrids and contact zones were strongly bimodal in all cases, but with measurable differences in levels of gene flow among contact zones. Using data available from previous studies on this complex and in a multivariate statistical framework, we compared the level of hybridisation in each contact zone with several predictor variables: nuclear and mitochondrial genetic divergence, distribution overlap, and morphological, ecological and physiological distances between species. Our results show that, despite the general trend in other taxa, the level of hybridization in Podarcis contact zones does not seem to be correlated to divergence time. We discuss how other mechanisms may have influenced this pattern.

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Niche divergence among closely related lineages can be informative on the ecological and evolutionary processes involved in differentiation, particularly in the case of cryptic species complexes. Here we compared phylogenetic relationships and niche similarity between pairs of lineages included in the Podarcis hispanicus complex to examine patterns of niche divergence and its role in the organization of current diversity patterns, as allopatric, parapatric, and sympatric lineages occur in the Western Mediterranean Basin. First, we used ecological niche models to characterize the realized climatic niche of each Podarcis hispanicus complex lineage based on topographic and climatic variables, to identify important variables, and to test for niche conservatism or divergence between pairs of lineages. Variables related to precipitation generally exhibited the highest contribution to niche models, highlighting the importance of rainfall levels in shaping distributions of Podarcis wall lizards. We found that most forms have significant differences in realized climatic niches that do not follow the pattern of mitochondrial divergence. These results lend support to the hypothesis that genetic divergence across Podarcis hispanicus complex most likely occurred in allopatric conditions, mostly with significant niche divergence. Competition after secondary contact is also suggested by the common occurrence of niche overlap between lineages that exhibit strictly parapatric distribution. The almost continuous distribution of Podarcis lizards in the study area appears to be a result of a combination of complementary suitable niches and competition, which seem two important mechanisms limiting geographic distributions and restricting the existence of extensive contact zones.

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This paper analyses the literature published in the last 17 years regarding records of albinism, axanthism, erythrism, leucism, melanism, nigrism or piebaldism, among other chromatic anomalies in Iberian herps.Whereas hipomelanistic disorders showed to be more widespread, melanism cases had a high occurrence pattern in the northern side of the Iberian Peninsula.We suggest that thermal melanism could be an adaptive advantage for ectotherms. Notwithstanding this, we also discuss the possible negative effect of the color aberrations taking into account differences in pre-dation pressure.

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Ecological factors are known to limit species geographical distribution. Lacertids, being ectotherms, are likely to be most influenced by thermal conditions but factors such as environmental humidity or species interactions may also be preponderant. Podarcis liolepis and P. muralis are lacertid species overlapping at a small scale in north-eastern Iberian Peninsula (IP). While P. liolepis ranges from the north-east of IP to southern France, P. muralis has a wide range across Europe but in IP is restricted to the north. Knowing and understanding the factors most likely to shape this pattern is the aim of this study. This has been assessed through three methodological approaches. Ecophysiological experiments in the laboratory were conducted to assess the thermal and hydric requirements of P. liolepis and P. muralis (manuscript I). In a sympatry area, field records of body temperature were taken and the influence of environmental factors on body temperatures were assessed (manuscript II). Correlative predictive models were performed in order to assess the putative current and future geographic distribution of both species (manuscript III). The ecophysiological data previously obtained was integrated in the interpretation of the modelling data. The three approaches revealed that environmental humidity and not temperature is the main abiotic factor constraining these species. However, evidence of possible interaction between both species was detected in terms of thermal physiology since the preferred body temperature of P. liolepis shifted upwards in the presence of P. muralis. Combining this information with the prediction for future scenarios of climate changes, it is expected that P. liolepis may be the most affected if climate gets to hot and dry since it would shift northwards and would disappear from some areas due to aridification. On the other hand, P. muralis’ range is likely to decrease but little fragmentation would occur.

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In ectotherms, environmental factors shape the distribution of species mediated by ecophysiological constraints such as thermal requirements and water stress. Species with different distributions along an environmental gradient are expected to show contrasting responses in thermal-gradient and water-stress lab experiments. We examined basic thermal and hydric physiological traits throughout the day in two related lizard species with different, but partially overlapping, distributions in the Iberian Peninsula: Podarcis liolepis (abundant but mostly restricted to northeastern Iberia) and P. muralis (restricted in Iberia but widespread across Europe). We expected P. liolepis to opt for higher preferred body temperatures and have lower water loss rates as compared to P. muralis. Surprisingly, results revealed no differences in preferred body temperatures between species or sexes. Conversely, interspecific differences in the temporal profiles of water loss were found. Results suggest that water availability rather than thermal environment shapes the biogeographical patters of both species.

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A critical review of the evolutionary biology of Iberian and North African lizard members of the genus Podarcis (Lacertidae) based on phylogeny, phylogeography, morphometrics, behavior, ecology and physiology is presented. The Iberomaghrebian region is inhabited by at least 12 different evolutionary lineages that group into a monophyl- etic clade (except Podarcis muralis). In contrast to the current taxonomy, the saxicolous ‘Podarcis hispanica’ is paraphyletic with respect to Podarcis bocagei and Podarcis carbonelli, two currently recognized species. Nodes in the phylogenetic tree are deep, resulting from old divergences, clearly preceding the Pleistocene. Nevertheless, more recent range changes as a result of glaciations are also evident. The most plausible evolutionary scenario for this group indicates both vicariant and dispersal events. Although parapatry between lineages is the rule, sympatry and even syntopy are frequent, but usually between ground-dwelling and saxicolous forms. Contacts between forms with similar habitat use are rare and local. Morphological distinctiveness between lineages has been demonstrated, indicating historical constraints. However, other characters have repeatedly evolved under similar environmental pressures independently of the evolutionary lineage. Strong sexual dimorphism derives from sexual selection and is attained before sexual maturity, although developmental restrictions exist. Variation between popu- lations is also important and derives from local variation in both natural (habitat, climate) and sexual (density) selective pressures. Evidence for short-term changes has already been found, particularly in insular populations. Reproductive isolation between syntopic forms and partner recognition are based on male–male competition and on visual and chemical recognition of females by conspecific males. Despite this ancient diversification, most forms maintain a degree of reproductive compatibility. Hybridisation may occur, but is limited, and there is evidence of selection against hybrids. The ecological analysis presented here does not support exploitative competition, but rather behavioral interference between forms. Ecomorph classification of lineages suggests character displacement between those with extensive range overlap. Finally, a critical assessment of the specific status of all lineages is provided and directions for further research are suggested.

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Podarcis hispanica is, in fact, a complex of several forms and ecophysiological traits previously investigated in one of them my be no longer generalisalbe to the others. Intraspecific variation of preferred temperatures (Tp) was analysed in a population of the NE form of P. Hispanica and compared with data on other forms coming from the literature. Adult lizards were collected in May, measured (SVL), weighed and exposed to a thermal gradient at nine time intervals. Tp changed with sex (males > females), size (positive correlation with body mass) and, more weakly, with time of day (early morning > mid morning and mid-day > afternoon). Two females decreased their Tp after egg-laying. Results corroborate previous studies demonstrating diel adaptability of Tp in lacertids and strong dependence on the individual’s condition. Tp should be interpreted as a compromise between different pressures including gonadal changes, embryo development and social behaviour. Furthermore, in a comparison restricted to males, the NE form attained lower Tp than other Iberian Podarcis not directly related in the phylogeny, which may indicate changes at evolutionary level.

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In the essay the finding and some morphological characteristics of a melanic example of Podarcis hispanica atrata found on the Columbretes archipelago (Castellón, Spain) is described.

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The incidence of cannibalism in the lizard Podareis hispaniea atrata in the Columbretes islands (Castelló de la Plana, Spain) under natural and captive conditions is described. Adult male lizards can ingest one hatchling in approximately four minutes. Hatchling digestability appeared to be very high.

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This study reports some observations on the predation 01 lizards (Podareis hispaniea atrata) by scorpions (Buthus oeeitanus), as well as on the predation 01 scorpions by this lizard species in the Columbretes islands (Castellion, Spain).

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This paper reports on the selection of substrate temperatures at oviposition sites by female lizards Podarcis hispanica atrata under laboratory conditions. Seven females deposited 11 clutches (a total of 34 eggs) in very small areas, representing 20 % of the available surface. The mean temperature of the sites used for egg laying was 25.7°C. Results are compared with incubation temperatures of the same species under natural conditions (Columbretes islands, Mediterranean, Spain).

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In this study we provide inlormation on the current status 01 two reptile species (Podareis hispanica and Tarentola mauritanica) in the Medes islands. Comparison 01 our data with published observations shows that the abundance 01 P. hispanica appears as substantially declined. We emphasize the importance 01 the conservation 01 biodiversity in small isolated islands and that the monitoring 01 the status 01 isolated populations is required to develop the appropriate management strategies.

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We studied aspects of the thermal biology and microhabitat selection of the endangered lizard Podarcis hispanica atrata during autumn in the field and laboratory. Body temperatures (T b ) of active lizards were within a narrow range, were largely independent of ambient temperatures, and exhibited little diel variation. Activity T b s largely coincided with the selected temperatures maintained in a laboratory thermogradient and with T b s that maximize running performance. Alternation of basking with other activities and shuttling between sun and shade were obvious aspects of thermoregulatory behaviour. Lizards shifted microhabitat use throughout the day. During early morning and late afternoon, basking lizards were restricted to rocky sites surrounded by shrubs. Near midday lizards used a wider array of microhabitats, and many moved in open grassy sites. Juveniles maintained lower activity T b s, had lower selected temperatures, and basked less frequently than the adults. Juveniles occupied open grassy patches more often than the adults. We discuss the relevance of our results for the conservation of this extremely rare lizard and the management of its habitats.

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The world range of the lizard Podarcis hispanica atrata is retricted to the Columbretes archipelago in the Mediterranean. We examined its actual distribution, abundance and some aspects of its natural history. Lizards were found in four islets (combined area c. 20 ha). Densities on the largest islet (13 ha) are high (> 600 lizards/ha), at least in favourable habitats. Population sizes on the smaller islets are extremely small (< 200 individuals). A considerable number of juvenile lizards was found in all populations. We report new information on body length distribution, sexual dimorphism, tail break frequency, escape behaviour, and diet composition. Recommendations for the conservation of this lizard and the improvement of its habitat are given and discussed in relation to landscape management programmes.

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We report reproductive characteristics (timing of sexual maturity, clutch frequency, clutch size, egg and hatchling sizes) of the Island Lizard Podarcis atrata. Reproductive performance was related to female size. The larger females initiate reproductive activities early in the season and produce two to four clutches annually, whereas the smaller individuals postpone reproduction until the season is well advanced and lay only a single clutch. Clutch size increased with female length and was slightly lower for clutches laid during June–July than for those produced during May. Among-clutch differences in egg mass were negatively correlated with clutch size, and this relationship is independent of female size. Larger clutches are also composed of less elongated (i.e., rounder) eggs than are smaller clutches, suggesting that the phenotypic trade-off between egg size and numbers is induced by limitations posed by the females` body volume, rather than by restrictions of energy availability. Comparison of reproductive traits of P. atrata to those of the closely related P. hispanica and P. bocagei indicates that the increase in body length during the evolutionary divergence of P. atrata was not associated with an increase in clutch size. This provides a clear exception to the general pattern found in lacertid lizards.

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Sequence data derived from a 306 bp fragment of mitochondrial cytochrome b and molecular variance estimates were used to investigate the genetic population structure of the endangered and endemic lizard Podarcis atrata of the Columbretes archipelago (Mediterranean, Spain). Our results show a very high and significant among-population genetic differentiation. FST values and phylogenetic analyses confirm the evolutionary distinctiveness of P. atrata populations, suggesting that the populations of these islands deserve special protection measures. The populations of the two islands Columbrete Grande and Mancolibre are less differentiated than those of Foradada and Lobo, and seem to have retained mainland haplotypes. This situation needs further attention as the origin of the mainland haplotypes is still unclear. If they are a result of recent introductions from mainland specimens, then they may represent a threat to the endemic lizards of the Columbretes islands.

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Endemic species and island populations are particularly prone to extinction. For this reason, their identification and protection is particularly important. Podarcis hispanica atrata is an endangered and endemic lacertid confined to the Columbretes Islands (19 ha, Mediterranean Sea, 39 degree 54`N, 0 degree 41`E, Spain). The status and nomenclature of P. h. atrata has a confused taxonomic history, but currently the taxon is considered as P. h. atrata. Populations of P. h. atrata differ from populations of P. h. hispanica in morphology, physiology, behavior, and life history. Because these differences may be ecogenetic, that is, habitat differences caused rapid adaptational change, this evidence alone is not sufficient enough to determine the taxonomic status of P. h. atrata. In this study, we used putatively neutral molecular markers (mitochondrial DNA sequences) to assess degree of divergence among populations of P. h. atrata relative to closely related taxa. The data are taxonomically relevant and provide a basis to develop conservation and management policies for this endangered lizard.

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In this study we present the first ecological data for the spiders Argiope lobata and A. bruennichi in Spain. We show data on density variation among two years (2004-2005), on the size of spiders and their webs, on sorne characteristics of the vegetation occupied by the two species, and on sorne aspects of their behaviour. The study has been conducted in the Columbretes islands (Mediterranean sea, Spain), taking the advantege of the recent colonization of the main island (Columbrete Grande, 13 ha) by the two spider species. We also discuss about the possible effect that such colonization could have on the endemic fauna of the island, particularly on the lizard Podareis atrata.

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Juveniles of a variety of lizard species exhibit bright tail colors that contrast clearly to the cryptically colored body. The `predator escape` hypothesis suggests that bright tail colors deflect the attack of a predator towards the lizard`s tail, which can be autotomised, and hence increase the probability of surviving a predatory attack. Although this hypothesis is widely accepted, surprisingly few empirical data exist to support it. Here, we examined the relative frequencies of predatory attack on plasticine replicas of juvenile lizards in the genus Podarcis with either green or brown tails in nine areas of Spain. Replicas with bright green tail color experienced, over a one-week period, the same rates of attack by birds and other predators as replicas with cryptic brown tails. However, the proportion of replicas that showed bill markings on the tail only was highest for the green-tailed replicas. In contrast, the frequency of predatory attacks towards the head or body was similar in the two groups of replicas. Our experiment appears to support the classical prediction of the adaptive value of a green autotomic tail in lizards. However, lizard species with multiple tail colors may also represent alternative anti-predator strategies that may have arisen by different selective pressures between habitats. We also suggest that the evolution of bright tail coloration in juveniles of Podarcis as an antipredator mechanism may be favored by the correlated evolution of morphological, physiological, and behavioral characteristics

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To understand the dynamics of small populations in arid environments, particularly in small insular ecosystems, it is crucial to understand the interactions between species inhabiting these environments. Here we investigate the interactions between lizards and scorpions in a small arid insular ecosystem in the Mediterranean. Experimental observations suggest that scorpions are regarded as prey by lizards. However, the response of lizards was sex-dependent, with males being more likely to attack scorpions than females. This can be associated with the larger body and head size, and bite forces in males, presumably making scorpions a profitable prey. Predation on lizards by scorpions has been also documented, suggesting a classic case of cross predation.

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This study reports on an experiment designed to quantify potential predation by birds on the lizard Podarcis hispanica atrata in the Columbretes islands (Castellón, Spain). We used plasticine replicas to examine predation risk of replicas as a function of proximity to vegetation and to potential predators, and to investigate the seasonal variation in patterns of predation. Predation on replicas was very high and intensive within only four days, and differed between study areas and seasons. Predation was more intense when replicas were closer to nests of gulls and far from vegetation. The Yellow-legged Gull (Larus cachinnans michaellis) was identified as the major species responsible for predation on lizard replicas.

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In this study we document the spatial and temporal variation in the density of two orb-web spiders (Argiope lobata and A. bruennichi) in the Natural Park of Columbretes islands (Castellón, Spain) during 3 con- secutive years (2004-2006). The masive presence of these species was detected for the first time in 2004 in the main island Columbrete Grande (13 ha). From then, we followed their population size. It is very important to document temporal fluctuations in the den- sity of invader species, mainly in small islands occupied by endemic vertebrate and invertebrate species that may suffer higher predation and competition for food.

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This paper reports the development of u new technique that may help to increase the number of lizards in areas where scarcity of oviposition sites constitutes the major limiting factor. We have created arttjicial egg-laying sites for the endangered subspecies of lizard Podarcis hispanica atrata in the Columbretes archipelago. Our results demonstrate that such urtiJcia1 sites were quick1y accepted bv the lizards us areas for basking und burrowing. Only one container was used for oviposition by one female, but the physical conditions in the artificial sites allowed successful development in 83’% of the eggs which were experimentally placed in them. Additional studies will be required to improve this technique and to determine if it might be applicable to other lizard species.

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We report the effects of incubation temperature on the duration of incubation and hatchling size in the insular lizard Podarcis hispanica atrata. Freshly-laid eggs were placed in artificial nests that were exposed to a natural cyclical variation of microclimate. We monitored substrate temperatures and egg size throughout the incubation period, and measured hatchling size at birth. The mean incubation period was 54 days at average incubation (substrate) temperatures that ranged from 23.5 to 26.5 C. We found a negative relation between the duration of incubation and mean incubation temperature. Overall survival of eggs to hatching was high (83%), even though eggs were frequently exposed for short periods (1-5 h) to high substrate temperatures (>35 C). We found no effect of duration of incubation or incubation temperature on the size of hatchlings.

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The lizard Podarcis atrata, endemic to the Columbretes archipelago (Mediterranean, Spain) occupies very small islands characterized by extreme aridity and a reduced availability of potential insect prey. The results of an experimental study have shown that adult lizards offered individuals of Ligia italica and Armadillo officinalis (marine and terrestrial isopods, respectively) consumed at high frequencies; with males consuming them at significantly higher frequencies than females. The results are discussed in light of the importance of the inclusion of marine prey into the diet in arid insular ecosystems.

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The updated distribution of the 40 amphibians and reptiles species recorded in Jaén province is showed off in 10x10 UTM squares. Distribution of 35 of them is enlarged with 323 new records, 152 for amphibians and 171 for reptiles. Main conservation problems detected for every species are expounded.

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The present article covers 20 years of field work by the authors. It provides data of 26 species of amphibians and reptiles, located in the southern sector of Catalonia (northeastern Spain), many of them in quadrants UTM 10 x 10, which had not been previously cited.

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Podarcis hispanica est signalé pour la première fois dans les départements du Tarn et de la Haute- aronne. Sur la base des données récoltées par les auteurs ou transmises par d`autres observateurs, les limites de la répartition connue de l`espèce dans les départements de l`Ariège, de l`Aude et des Pyrénées- Orientales sont aussi présentées.

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Comparative analyses were carried out about the variability of the pileal shields of different species belonging to the family Lacertidae. The results of the comparisons have been evaluated in respect to the phylogenetical relationships existing between the different genera and species of the family.

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We studied the effect of habitat fragmentation on female reproductive investment in a widespread lacertid lizard ( Psammodromus algirus) in a mixed-forest archipelago of deciduous and evergreen oak woods in northern Spain.We captured gravid females in fragments (≤10 ha) and forests (≥ 200 ha) and brought them to the laboratory, where they laid their eggs. We incubated the eggs and released the first cohort of juveniles into the wild to monitor their survival. Females from fragments produced a smaller clutch mass and laid fewer eggs (relative to mean egg mass) than females of similar body size from forests. Lizards did not trade larger clutches for larger offspring, however, because females from fragments did not lay larger eggs (relative to their number) than females from forests. Among the first cohort of juveniles, larger egg mass and body size increased the probability of recapture the next year. Thus, fragmentation decreased the relative fecundity of lizards without increasing the quality of their offspring. Reduced energy availability, increased predation risk, and demographic stochasticity could decrease the fitness of lizards in fragmented habitats, which could contribute to the regional scarcity of this species in agricultural areas sprinkled with small patches of otherwise suitable forest. Our results showthat predictable reduction of reproductive output with decreasing size of habitat patches can be added to the already known processes that cause inverse density dependence at low population numbers.

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Background Lizards of the genus Podarcis are widespread in the Mediterranean region, including islands and island archipelagos. These small-bodied lizards have a predominantly protective green-brown colouration. However, some populations display unusual patterns, in which the colouration is predominantly blue or uniformly black. This study explores the factors that influence this chromatic variation, whether environmental (climate and island conditions) or evolutionary (phylogenetic trait conservatism). The colouration of 1400 individuals (27 species) was analysed in the CIELAB colour space. Results Pagel’s λ indicated that colouration is weakly conserved within phylogenetic lineages. Although the island surface plays a key role in the chromatic variability of these lacertids, geographic isolation and climate hold less influence. The colouration of some small island populations tends to be uniform and dark, possibly due to intense intraspecific competition and lower predatory pressure. Conclusions This study highlights the importance of island populations in understanding the processes that favour the emergence of extreme phenotypes in small ectothermic vertebrates.

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Mammals and birds are capable of navigating to a goal using learned map-like representations of space (i.e. place learning), but research assessing this navigational strategy in reptiles has produced inconclusive results, in part due to the use of procedures that do not take account of the peculiarities of reptilian behavior and physiology. Here I present a procedure suitable for testing spatial cognition that exploits a naturally evolved, ethologically relevant ability common to many lizards (i.e. refuge seeking behavior). The procedure requires lizards to learn the location of an open refuge inside a rectangular arena containing artificial refuges in every corner, using distal extramaze visual cues and with no local cues marking the location of the open refuge. The procedure probes the lizards’ place learning ability and effectively rules out the use of egocentric and response-based strategies. The described procedure was successfully used to demonstrate place learning in a lacertid lizard (Podarcis liolepis). Over the course of two weeks of training both the latency to entering the open refuge and the number of corners visited in each trial decreased gradually, indicating that learning had taken place in over 60% of the lizards tested. These results confirm that, under certain circumstances, lizards are capable of navigating to a goal using a place learning strategy.

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The introduction of alien species to islands by human activity can cause catastrophic consequences for small populations of island endemics. Hybridization between the endangered and endemic insular lizard Podarcis atrata (Boscá, 1916) from the Columbretes Islands (Mediterranean, Spain) and the common mainland lizard Podarcis hispanica (Steindachner, 1870) could potentially occur because mainland haplotypes have already been detected in the islands, the two species are closely genetically related, and the frequency of visitors to these islands is increasing. However, reproductive decisions of lizards are often mediated by species recognition mechanisms based on chemical cues. On the basis of this observation, even if some mainland P. hispanica lizards were introduced to the islands, interspecific recognition might make rare an eventual hybridization with the insular P. atrata. We examined interspecific chemical recognition between the insular P. atrata and the mainland P. hispanica. Our results showed that lizards of both sexes responded more strongly (i.e., directed a significantly higher number of tongue flicks) to scents of conspecific individuals than to scents of heterospecifics. Chemical recognition of conspecifics by endemic island P. atrata lizards may reduce the occurrence of hybridization with introduced mainland P. hispanica lizards and protect the insular gene pool.

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Chemical signals can be the basis of interspecific recognition and speciation in many animals. The Columbretes Islands wall lizard, Podarcis atrata is very close genetically to the mainland Iberian wall lizard Podarcis hispanica. However, a previous study suggested that chemosensory interspecific recognition would avoid reproductive interactions and hybridization between these two species. These results suggested that chemicals used in intraspecific communication might differ in composition and/or proportions between these two species. In this paper, we used gas chromatography–mass spectrometry (GC–MS) to characterize the chemical composition of the lipophilic fraction from femoral gland secretions of male P. atrata and P. hispanica. The analysis showed that chemicals found in femoral secretions varied in composition and proportions between species and between populations. Seven steroids and two unidentified waxy esters, were exclusive of P. atrata lizards from the islands. In contrast, nine steroids and other six compounds were only found in mainland P. hispanica. There were also differences in proportions of shared compounds between species. Moreover, all these differences were higher between P. atrata and P. hispanica than between any population of P. hispanica. Chemical differences might be consequence of genetic differences, but they could also be explained by adaptation to different habitats with different climatic conditions or diet resources. Compounds that are specific of each species, or differences in the pattern of compounds, could explain species recognition. Therefore, these results of chemical composition and previous studies of chemosensory recognition reinforce the fact that the genetic differences between P. hispanica and P. atrata may result in an effective reproductive isolation between these two taxa.

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Major histocompatibility complex (MHC) genes are crucial for the adaptive immune response of jawed vertebrates. Their variation, reaching extreme levels, is driven mainly by an arms race between hosts and pathogens. One hypothesised mechanism contributing to MHC polymorphism is adaptive introgression, the exchange of genetic variants between hybridising species favoured by selection, yet its effect on MHC variation is poorly understood. Detection of adaptive MHC introgression, though challenging, may be facilitated by the analysis of species complexes forming multiple hybrid zones. Here, we investigated MHC introgression in six hybrid zones formed by seven species of Podarcis lizards inhabiting the Iberian Peninsula. To differentiate adaptive introgression from neutral introgression, we compared the patterns of gene exchange in MHC and genome-wide markers. We found elevated sharing of MHC alleles in the proximity of contact beyond the areas of detectable genome-wide admixture in most hybrid zones and, in half of them, asymmetric MHC exchange. In general, the elevated MHC allele sharing between species pairs with abutting ranges compared to geographically isolated species pairs also supports the prevalence of introgression. Collectively, our results demonstrate widespread MHC introgression in the Iberian Podarcis complex and suggest its adaptiveness. Contrary to previous results from Triturus newts, we did not observe differences in the rate of introgression between MHC classes. Our work adds support to the emerging view of adaptive introgression as a key mechanism shaping MHC diversity. It also raises questions about the effect of elevated MHC variation and factors leading to the asymmetry of adaptive introgression.

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The frequent lack of biodiversity data on municipal areas favoured the performance of a herpetological study to determine the number of herpetological species in the four most significant types of habitat within the municipality of Navaridas (province of Alava). A number of different sampling surveys were performed (transects, and water net points) during 2015 and 2016. 15 herpetological species (8 amphibians and 7 reptiles) were found. Protected areas showed higher diversity levels, especially in the lake and associated holm oak woodland of Navaridas. The results suggest the possible benefit of low stone walls between vineyard for Podarcis liolepis. This study establishes a starting point for further research projects based on the population densities and evolution of herpetological species, and an assessment of the possible effects of agricultural activities on these populations.

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Climatic conditions changing over time and space shape the evolution of organisms at multiple levels, including temperate lizards in the family Lacertidae. Here we reconstruct a dated phylogenetic tree of 262 lacertid species based on a supermatrix relying on novel phylogenomic datasets and fossil calibrations. Diversification of lacertids was accompanied by an increasing disparity among occupied bioclimatic niches, especially in the last 10 Ma, during a period of progressive global cooling. Temperate species also underwent a genome- wide slowdown in molecular substitution rates compared to tropical and desert-adapted lacertids. Evaporative water loss and preferred temperature are correlated with bioclimatic parameters, indicating physiological adaptations to climate. Tropical, but also some popu- lations of cool-adapted species experience maximum temperatures close to their preferred temperatures. We hypothesize these species-specific physiological preferences may con- stitute a handicap to prevail under rapid global warming, and contribute to explaining local lizard extinctions in cool and humid climates.

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Recent genetic works have suggested that the Iberian wall lizard Podarcis hispanicus (Steindachner, 1870) sensu lato may in fact be a species complex, yet many of its taxa have not been formally described. We redefine the nominotypical Podarcis [hispanicus] hispanicus sensu stricto using univariate and multivariate morphological analyses and pinpoint its geographical distribution. Podarcis [hispanicus] hispanicus is a small wall lizard characterized by a flattened head and body, the general lack of a masseteric shield, a striped dorsal pattern, the frequent bifurcation of the anterior part of the vertebral strip, the belly almost always white and the tail intense blue in young specimens. Its distribution is restricted to the Spanish Levant (provinces of Alicante, Murcia and Almeria).

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A new population of Podarcis liolepis has been discovered in the suburb of Toulouse (France), corresponding to the third known place of this species for the province of Haute-Garonne and the first of the nominotypic subspecies for the region of Midi-Pyrénées. Origin of this population is discussed and morphological traits to identify the nominotypic subspecies are described.

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Podarcis liolepis was formally considered as an Iberic element recently arrived in France, reaching the western Rhône river border but no trespassing it. The discovery of two new stations East of the Rhône River represent the first record for the Provence, in two new French departments, Drôme and Vaucluse.

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La observación de ejemplares de P. muralis y P. hispanica en la línea de costa que contacta con el mar, ha permitido obtener información más amplia sobre su distribución en el área septentrional más extrema, así como del di- ferente uso que hacen del nicho espacial. Ambas se revelan, una vez más, como especies antropófilas, apareciendo preferentemente P. muralis sobre acantilados de sustrato calizo y suelo cubierto de vegetación, frente a P. his- panica, que prefiere el sustrato areniscoso y suelos más despejados. Los datos conocidos sobre su distribución, junto con los aquí aportados, permiten esbozar un esquema biogeográfico, en el que lo más destacable se- ría la penetración que, a través de ciertos valles térmicos interiores, ha se- guido P. hispanica -junto con otras formas termófilas de la herpetofauna- , para colonizar el área costera, donde la suavidad de las condiciones climáti- cas ha facilitado su asentamiento.

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Residual yolk is assumed to be an important source of energy and nutrients during early life in nonmammalian amniotes. Available data show that the mean size of residual yolk is far smaller in lizards than in turtles, snakes, crocodiles, and birds, raising a question of whether residual yolk is of functional significance in lizards. Here, we compared data from 26 lizard species with those from other nonmammalian amniotes to test the hypothesis that residual yolk is functionally less significant in species producing more fully developed offspring. In our sample, species mean offspring water contents ranged from 73% to 84% of body wet mass; species mean proportions of carcass dry mass, fat-body dry mass, and residual yolk dry mass to offspring dry mass ranged from 84% to 99%, 0% to 5.0%, and 0% to 14.4%, respectively. Lizards are, on average, more fully developed at hatching or birth than snakes, as revealed by the fact that the mean proportion of carcass dry mass to body dry mass and offspring water contents were both higher in lizards than in snakes. We conclude that the functional significance of residual yolk during early life is generally less evident in lizards. Even in the lizards where residual yolk is of potential functional significance, this portion of yolk contributes little, if any, to postembryonic growth. Future work could usefully collect data across a wider spectrum of reptile taxa to establish a precocial–altricial continuum and test the hypothesis that species with a smaller amount of residual yolk are closer to the precocial end of the continuum.

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In recent years there has been an exponential increase in the amount of published DNA sequences, as it has become one of the most popular sources of data for reconstructing phylogenies. A number of studies have attempted to resolve the relationships of groups of lacertids using mitochondrial DNA sequence data, in particular Podarcis and Gallotia (e.g. THORPE et al., 1994; 1996; GONZALEZ et al., 1996; FU et al., 1997; HARRIS, 1997; HARRIS et al., 1998a; 1999; CASTILLA et al., 1999a,b). Reassessment of these data suggest some alternative phylogenies to those previously published, and also highlight possible problems. Two studies have attempted to unravel the phylogeny of the whole family using mitochondrial DNA sequences (FU, 1998; HARRIS et al., 1998b). By combining these essentially congruent data sets, and reanalysing levels of support for the phylogenies produced, a more complete estimate of relationships across the family is produced. This is then compared to alternative sources of evidence, primarily morphological (ARNOLD, 1989), biochemical (MAYER & BENYR, 1994) and karyological (OLMO et al., 1993). Reasons for discrepencies are discussed, as are the relative advantages and disadvantages of the different sources of phylogenetic inference.

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Aim To examine the effect of a known geological barrier on genetic variation within a wall lizard species complex. Location The Iberian Peninsula and North Africa. Methods Sequencing of partial 12S rRNA and cytochrome b mtDNA. Results The current distribution of genetic variability is not related to the opening of the Strait of Gibraltar. Conclusions Podarcis hispanica in North Africa is probably a species complex. The Strait of Gibraltar should not be used as a known barrier to gene flow in other land based organisms without careful sampling to test for multiple crossings since its formation.

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Phylogenetic relationships between species and morphotypes of Podarcis wall lizards from the Iberian Peninsula and north Africa were estimated using partial 12S rRNA and cytochrome b mitochondrial DNA sequences. All species except Podarcis hispanica form monophyletic units. P. hispanica is paraphyletic, although all identified morphotypes are monophyletic. These morphotypes represent highly divergent lineages showing 10–15% pairwise sequence divergence with the cytochrome b gene. The data suggest that P. hispanica is a species complex. We recommend using P. hispanica* until additional sampling delimits the number and ranges of species currently referred to P. hispanica. P. carbonelli, which has recently been raised to species status, is confirmed as a genetically distinct form. P. atrata is genetically distinct, but much more closely related to some populations of P. hispanica than previously thought.

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1. Research addressing the effects of global warming on the distribution and persistence of species generally assumes that population variation in thermal tolerance is spatially constant or overridden by interspecific variation. Typically, this rationale is implicit in sourcing one critical thermal maximum (CTmax) population estimate per species to model spatiotemporal cross‐taxa variation in heat tolerance. Theory suggests that such an approach could result in biased or imprecise estimates and forecasts of impact from climate warming, but limited empirical evidence in support of those expectations exists. 2. We experimentally quantify the magnitude of intraspecific variation in CTmax among lizard populations, and the extent to which incorporating such variability can alter estimates of climate impact through a biophysical model. To do so, we measured CTmax from 59 populations of 15 Iberian lizard species (304 individuals). 3. The overall median CTmax across all individuals from all species was 42.8 °C and ranged from 40.5 to 48.3 °C, with species medians decreasing through xeric, climate‐generalist and mesic taxa. We found strong statistical support for intraspecific differentiation in CTmax by up to a median of 3 °C among populations. We show that annual restricted activity (operative temperature > CTmax) over the Iberian distribution of our study species differs by a median of > 80 hours per 25‐km2 grid cell based on different population‐level CTmax estimates. This discrepancy leads to predictions of spatial variation in annual restricted activity to change by more than 20 days for six of the study species. 4. Considering that during restriction periods, reptiles should be unable to feed and reproduce, current projections of climate‐change impacts on the fitness of ectotherm fauna could be under‐ or over‐estimated depending on which population is chosen to represent the physiological spectra of the species in question. Mapping heat tolerance over the full geographical ranges of single species is thus critical to address cross‐taxa patterns and drivers of heat tolerance in a biologically comprehensive way.

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Sexual dimorphism of relative head size is a widespread phenomenon in lizards, males having larger head/trunk ratios than females. In an attempt to explain this sexual dimorphism several hypotheses have been formulated. The two most frequently cited ones are: 1) sexual selection acting on those structures important in intrasexual competition and 2) natural selection for reduction of food competition between the sexes. In the insular subspecies of Podarcis hispanica (P h. atrata) males tend to have significantly larger heads than similarly sized females. We here test an implicit assumption of the dietary divergence hypothesis, namely that an increase in head size results in an increase in gape width and/or bite force, thereby allowing the larger headed sex to exploit larger prey classes. Using a static bite force model, we calculated the magnitude of bite forces for given directions at given positions on the jaws and for different head sizes. We experimentally determined the hardness of three different prey items and compared the data to the maximal bite force produced by both sexes. Our results suggest an important difference in male and female bite capacity, which may bear significant ecological relevance, and are in agreement with the implicit assumption of the dietary divergence theory.

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Presentamos nuevos registros de distribución en malla UTM de 10x10 km para Psammodromus algirus, Coronella austriaca y Vipera latastei en Soria, Castilla y León (España). Así mismo reportamos la sintopía de Podarcis muralis y Podarcis liolepis en dicha área. Adicionalmente, proveemos información detallada de la ocurrencia de 17 especies de reptiles y anfibios como contribución al esfuerzo de elaboración de mapas de herpetofauna de Soria.

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Lizards often respond to predators by hiding in sunless refuges, but this eliminates opportunities for thermoregulatory basking. Hiding can therefore lower body condition. Furthermore, in ectotherms basking is important to induce fever and activate an immune response. A potential trade-off therefore exists between lowering predation risk and elevating body temperature to fight infection. Such a trade-off could be habitat dependent if habitats differ in the relative risk of predation versus that of acquiring or countering an infection. Here we take an experimental approach to test whether lizard basking behavior is affected by a trade-off between predator avoidance and fighting an infection. We quantified the anti-predator behavior of male lizards (Podarcis liolepis) both before and after they were immune challenged (injected with LPS) or not (injected with PBS control). To test the generality of any trade-off, we tested lizards from both an urban and a natural habitat. We found that males spent less time hiding following a simulated predator attack after they had been immune challenged than before, but this decline was only significant for males from the natural habitat. We also tested whether morphological traits, body condition, and immune response level explained variation in male hiding time. In the natural habitat, but not in the urban habitat, males with relatively small heads hid for significantly longer. In conclusion, we show that lizard anti-predator behavior is affected by an immune challenge. Habitat differences in the factors that predict hiding time offers potential insights into why this might be the case.

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Cryptic species complexes represent groups that have been classified as a single species, because of the difficulty in distinguishing its members morphologically. Morphological investigation following the discovery of cryptic diversity is crucial for describing and conserving biodiversity. Here we present a detailed account of morphological variation in a group of Iberian and North African Podarcis wall lizards of the family Lacertidae, trying to elucidate the morphological patterns observed between known mitochondrial lineages. Our results reveal very high morphological variation within lineages, considering both biometric and pholidotic traits, but also indicate that lineages are significantly different from each other. The main sources of variation, both globally and between lineages, arise from body size, head dimensions, and limb length, possibly pointing to underlying ecological mechanisms. A combination of body size, body shape, and continuous pholidotic traits allows a relatively good discrimination between groups, especially when comparing one group with the rest or pairs of groups. However, ranges of variation greatly overlap between groups, thereby not allowing the establishment of diagnostic traits. The high morphological variation observed indicates that external morphology is not particularly useful for species delimitation in this group of lizards, as local adaptation seems to play a major role in within- and between-group differentiation.

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Size and shape sexual dimorphism have long been of great interest for animal biologists and herpetologists. Although general trends do exist, every species may represent a particular case and intraspecific variation in the degree and patterns of sexual dimorphism is not rare. In the case of P. hispanica, which presents a high morphological variability and whose intraspecific taxonomy is yet not well defined, interpopulations differences could help to elucidate the morphological patterns observed. The results of a morphological analysis of a population belonging the NE form of P. hispanica are here provided. A marked sexual dimorphism exists for the characters studied, both biometric and pholidotic, although sexual dimorphism in SVL is absent. Sexual differences in pholidotic characters, rarely examined, were marked, not only in femoral pores and ventral scales which are the characters usually studied in lizards. But also in gular scales and subdigital lamellae. The patterns of size and shape sexual dimorphism were in some cases also refelcted in the analysis of static allometries. Influences of sexual and natural selection on those traits are discussed.

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Evolutionary inference based on molecular phylogenetic methods has profoundly modified the way that we understand biological diversity, unravelling a higher evolutionary diversity than previously considered. An exemplary case of this is the group of Iberian and North African Podarcis wall lizards. More investigated than any other reptile group in Europe, the Podarcis hispanica species complex comprises unexpectedly high levels of phylogenetic diversity and illustrates how the discovery of further cryptic diversity may entangle evolutionary inference. In the present study, we report on the discovery of two new mitochondrial lineages in this species complex, reassess the phylogeny of the group, infer the age of major phylogenetic splits, and provide a detailed description of the geographical distributions of all known mitochondrial DNA lineages. Our data show that the differentiation of major lineages is older than previously considered, in most cases predating the Messinian salinity crisis. The new lineages discovered and their position in the phylogeny of the group profoundly modify previous biogeographical scenarios, clearly showing that the area today corresponding to the south-eastern corner of the Iberian Peninsula is a very important centre of diversification. The dating obtained for the differentiation of the lineages currently inhabiting this area coincides with the complex geological events that took place during the Miocene/Pleistocene transition, supporting the idea that both land movements and dramatic climatic oscillations during that period could be involved. Finally, the discovery of these new lineages, together with the observed distribution patterns, not only further augments the uncertainty associated to our understanding of the evolutionary history of this group of lizards, but also points to new areas of interest for future investigation.

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Psammodromus algirus, Podarcis hispanica, Timon lepidus and Tarentola mauritanica were observed in the surroundings of Lloret de Mar, NE Spain.

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On our return trip from the 2014 meeting of the AG Lacertiden, we observed fifity Podarcis muralis, four P. liolepis and one Lacerta agilis at the Burg Hardenberg, Lower Saxony. The ratio of observed P. liolepis seems to have decreased compared to 2011.

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Environmental enrichment has proven to be a useful and effective welfare tool in order to evaluate and enhance the well‐being of captive animals, but only when it is based on detailed knowledge of each species` natural behaviour. Chemoreception is fundamental to many aspects of reptilian biology; however, sensory enrichment with chemical stimuli has rarely been applied to reptiles. In this study, we evaluate the use of chemosensory enrichment as a method to enhance the welfare of Podarcis liolepis. For seven weeks, we exposed field‐caught males to scents from donor conspecific males collected on pieces of filter paper (i.e., “enriched” group, n = 18), and compared their behaviour to that of control males provided with unscented pieces of filter paper (n = 18). We measured the occurrence of normal (e.g., locomotion) and abnormal (escape attempts) behaviours each day for three weeks. In addition, we conducted two exploration tests and a visual barrier test. Compared to controls, enriched lizards showed a consistent long‐term decrease (29%–38%) in the occurrence of escape attempts. During exploration tests, enriched lizards spent less time performing escape attempts and devoted more time to perching than controls. As expected, both control and enriched lizards showed a reduction of time in locomotion and an increase in the time spent perching between the first and second exploration test, but these changes were significantly more pronounced for enriched animals. Taken together, our results suggest improved welfare of enriched animals, as they spent less time engaging in abnormal behaviours, more time in normal behaviours, and showed signs of faster habituation to a novel environment. Chemosensory enrichment is a relatively simple enrichment strategy that could potentially be applied to improve the welfare of a wide range of captive lizards, and reptiles at large.

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Identifying species is an important task in biology, medicine, pharmacology, agriculture and biodiversity conservation. Modern taxonomy, the discipline devoted to the description and identification of species, faces a decrease in its workforce while dealing with the challenge of describing the Earth’s vanishing biodiversity. In this context, automated identification of species becomes of major importance. Automated image identification is a thriving field of machine learning, with deep learning algorithms based on convolutional neural networks revolutionizing the field in recent years. These methods are also taking their first steps in the identification of images of biological taxa in a variety of different contexts and taxonomic scopes. In this work, we took a deep learning approach to classify images of Iberian and North African wall lizards, a group of cryptic species in which identification requires expert intervention. We addressed two problems: 1) a two-class problem focusing on the distinction between two species, Podarcis bocagei and Podarcis lusitanicus and 2) a nine-class problem involving all the species currently described in the group. Three different deep learning architectures were tested in both cases. In the two-class problem, classification success was high, reaching as high as 97.1% and 95.9% for ensemble models applied to male and female lizards, respectively. Classification in the nine-class problem was not as successful, highlighting the difficulties inherent to this group of cryptic species. However, results improved when predictions from different perspectives were combined, reaching 95.3% and 89.7% for males and females, respectively. These results suggest the utility of deep learning algorithms in the identification of cryptic species, providing promising resources in the taxonomical, evolutionary and conservation research.

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Chemosystematic investigations on the position of Lacerta jayakari and the genera Gallotia and Psammodromus (Sauria; Lacertidae) 13 species of the genera Lacerta, Psammodromus and Gallotia (L. erhardii, L. filfolensis, L. bedriagae, L. cappadocica, L. jayakari, L. vivipara, L. trilineata, L. agilis, P. hispanicus, P. algirus, G. atlantica, G. galloti, G. stehlin:) have been examined by means of micro complement fixation tests (albumin) as well as by proteinelectrophoresis concerning 14 genetic loci. The results and their con-sequences are the follows: 1. L. cappadocica, L. bedriagae, and L. erhardii/L. filfolensis are representatives of phylogeneticly equivalent groups (subgenera). Their isolation in the Middle Miocene corresponds with the first contacts of the landmasses of Eurasia and Africa. 2. Lacerta jayakari may be considered as a representative of a further group equivalent in this aspect; though the immunological results would not exclude a position near the green lizards (subgenus Lacerta s. str.). 3. Because of this isolated position of L. jayakari for this species (and Lacerta cyanura) the subgenus Omanosaura subgen. nov. has been established. 4. Psammodromus hispanicus and P. algirus have probably been isolated of each other since the Middle Miocene, too. 5. The radiation of the genus Gallotia corresponds closely with genesis of the Canary Isles.

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Investigated the albumins from 31 species and subspecies of the genus Lacerta s.1. by means of the immunoloical techniue Micro-Complement-fixation. Dendrograms were computed by means of the ktch-MargJiash algorithm. Moreover the taxa of the Lacerta saxicola and the Lacerta danfordi comlexes were analyzed by electrophoresis concerning 14 genetic loci. The results were discussel together with karyological and genitalmorphological data so far known yielding the following relationships: Lacerta graeca shows a closer relationshi to the subgenus Podarcis as does the quite closely related species air L. dugesii and L. perspicikata. Lacerta Iaevis probabl is connected closely to Podarcis, too. Lacerta parva shows a closer relationship to the `small izards` than to the sub-genus Lacerta s. str. A clear decision concerning the position of the Lacerta danfordi complex according to chemosystematical data was not possible. It is isolated from the other species probably about as long as L. puma. Biochemical differences between its `species` are too small to confirm their taxonomical revalorisation by Eiselt and Schmidtler Within the Lacerta saxicola complex the investigated taxa show a close relationship. Lacerta praticola and L. derjugini were definitly assigned to this group.

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Aim Temperature influences most components of animal ecology and life history – but what kind of temperature? Physiologists usually examine the influence of body temperatures, while biogeographers and macroecologists tend to focus on environmental temperatures. We aim to examine the relationship between these two measures, to determine the factors that affect lizard body temperatures and to test the effect of both temperature measures on lizard life history. Location World-wide. Methods We used a large (861 species) global dataset of lizard body temperatures, and the mean annual temperatures across their geographic ranges to examine the relationships between body and mean annual temperatures.We then examined factors influencing body temperatures, and tested for the influence of both on ecological and life-history traits while accounting for the influence of shared ancestry. Results Body temperatures and mean annual temperatures are uncorrelated. However, accounting for activity time (nocturnal species have low body temperatures), use of space (fossorial and semi-aquatic species are ‘colder’), insularity (mainland species are ‘hotter’) and phylogeny, the two temperatures are positively correlated. High body temperatures are only associated with larger hatchlings and increased rates of biomass production. Annual temperatures are positively correlated with clutch frequency and annual longevity, and negatively correlated with clutch size, age at first reproduction and longevity. Main conclusions Lizards with low body temperatures do not seem to have ‘slower’ life-history attributes than species with high body temperatures. The longer seasons prevalent in warm regions, and physiological processes that operate while lizards are inactive (but warm enough), make environmental temperatures better predictors of lizard life-history variation than body temperatures. This surprisingly greater effect of environmental temperatures on lizard life histories hints that global warming may have a profound influence on lizard ecology and evolution.

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Le Lézard catalan Podarcis liolepis a fait l`objet de deux observations récentes à des altitudes élevées (1850 m et 2750 m) dans le département de l`Ariège (France). L`altitude de ces deux observations est supérieure à l`altitude maximale précédemment connue chez l`espèce en France (1651 m) et l`une d`elles s`avère même supérieure au précédent record d`altitude enregistré chez P. liolepis pour l`ensemble de la chaîne pyrénéenne (2650 m).

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In this contribution we report the first case of albinism in Catalonian wall lizard (Podarcis liolepis) throughout its distribution, localized in Cànoves, Vallès Oriental (Catalonia).

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In the municipality of Gava, there is currently a total of 25 species of amphibians and reptiles (one of the order Urodela, eight of the order Anura, two of the genus Chelonia, seven of the clade Sauria and seven of the suborder Serpentes). Two of the species – Trachemys scripta and Discoglossus pictus – were introduced in recent times. Worthy of note is the presence of the last-known population of the Western Spadefoot (Pelobates cultripes) in the municipalities of the deltaic plain, and the location of the last population of the Natterjack Toad (Bufo calamita) on the same plain. The location of the Discoglossus pictus in the municipality of Gava is indicative of the expansion of this species, which was introduced in 2004 into the municipality of el Prat de Llobregat.

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Anhand morphologischer Merkmale wird die Verbreitungsgrenze von Podarcis hispanicus (s.str.) und Podarcis liolepis in der Provinz Alicante (Spanien) diskutiert. Die Auswertung unserer Ergebnisse stützt die These, dass es sich hier um geografisch klar abgegrenzte Arten mit nur wenigen Überlappungen handelt. Die bisher bekannte nördliche Verbreitungsgrenze von Podarcis hispanicus (s.str.) liegt bei der Stadt Elche. Nach unseren Erkenntnissen erstreckt sich das Verbreitungsgebiet jedoch 50 km weiter nach Nordosten. Die Verbreitungsgrenze korreliert außerdem mit einer klimatischen Grenze.

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Der Artikel berichtet über eine bislang unbeschriebene „gestreifte Form“ von Podarcis liolepis im Süden Spaniens und diskutiert deren taxonomischen Status. Anschließend werden Rückschlüsse auf das Verbreitungsgebiet von Podarcis liolepis in Südost-Spanien gezogen und Unterscheidungskriterien zu den benachbarten Arten beschrieben.

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Aim We discuss biogeographical hypotheses for the Mediterranean lizard species Podarcis and Teira within a phylogenetic framework based on partial mitochondrial DNA sequences. Methods We derived the most likely phylogenetic hypothesis from our data set (597 aligned positions from the 12S rDNA and phenyl tRNA) under parsimony, distance and maximum likelihood assumptions. Results The species usually included in Teira do not form a strongly monophyletic clade. In contrast, the monophyly of the genus Podarcis is rather well supported. Seven lineages are identified in the genus; in order of appearance within the tree, these are: the Balearic pityusensis and lilfordi pair, the sicula complex, a Tyrrhenian tiliguerta and raffonei pair, muralis, the Siculo-Maltese filfolensis and wagleriana pair, the Balkan group (erhardi, peloponnesiaca, milensis, melisellensis and taurica), and the Ibero-Maghrebian group (bocagei, atrata, hispanica and vaucheri). Conclusions The origin of the three European genera of lacertid assayed (Lacerta, Teira and Podarcis) is hypothesized to have occurred in the Oligocene. For Podarcis, a possible scenario of a Miocene diversification is derived from the sequence data, and the zoogeography of the lineages are discussed in relation to the palaeogeography of the Mediterranean. It is hypothesized that in the early history of the genus the main lineages separated by rapid, numerous and close events that produced a starting point very similar to a polytomy, hard to resolve by parsimony analysis of the data set.

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Widespread species often show extensive phenotypic variation due to the contrasting abiotic and biotic factors that shape selective pressures in different environments. In this context, the gradual and predictable patterns of variation in climatic and environmental conditions found in mountain areas offer a great opportunity to explore intraspecific phenotypic variation. For instance, temperature is negatively correlated with altitude and virtually all aspects of the behavior and physiology of ectotherms are sensitive to body temperature. In this work, we tested the hypothesis that morphology, dorsal and ventral coloration and the chemical profile of femoral secretions show interpopulational and seasonal variation in the lacertid lizard (Podarcis liolepis). We compared lizards from three populations inhabiting lowland and highland habitats in the French Pyrenees that were closely related genetically. We found that highland lizards were larger, stockier, had longer heads and more femoral pores and had a darker dorsal coloration than lowland ones. In addition, we detected interpopulational differences in both the abundance and the richness of chemical compounds in the glandular secretions, and we also found seasonal variation in the overall chemical composition. Dorsal and ventral coloration differed seasonally and between populations. Ventral and dorsal brightness were higher in lowland than in highland lizards in the reproductive season whereas the reversed trend was found in the non-reproductive season but only for dorsal brightness. In addition, all lizards had browner dorsal coloration in the non-reproductive season, and lowland lizards were greener in the reproductive season. By integrating information from both visual and chemical systems, our works offers a comprehensive view of how these lizards communicate in a multimodal context.

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Among the Lacertidae, the Catalan Lizard (Podarcis liolepis cebennensis) is rather scarce and unobtrusive. It is often goes unnoticed. The regional literature details the fact that this lizard occupies natural, sunny locations. Surveys carried out over recent years have shown, however, that it can be found in directly man-made sites such as walls. This species can be confused with the Wall Lizard (Podarcis muralis) and digital photography is an invaluable tool to help specialists in confirming records. The two species can be found in the same habitats, hence it is imperative to avoid misidentification in order to monitor changes in the population of these species.

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Evolution in islands and islets often leads to changes in chromatic characters of animals. In many cases, the result of these changes is an extravagant pattern, the loss of some chromatic character and/or the loss of its function, or the fixation of rare phenotypes. These changes are often explained by historical events as well as by several selective processes. The coloration of Podarcis lizards provides many examples of insularity driving changes in coloration. Podarcis liolepis atrata, an insular clade that inhabits the Columbretes archipelago, (a group of small volcanic islets Mediterranean coast) shows some differences respect to the general pattern found in continental P. liolepis populations. However, no study has addressed these differences objectively (i.e. using reflectance spectrophotometry techniques). Here, we compare the coloration of P. liolepis atrata with those shown by a continental population of P. liolepis liolepis (Penyagolosa massif; Iberian System), focusing on male coloration. Although data from other populations may be necessary for understanding the evolution of P. liolepis coloration, our spectral analyses reveal chromatic differences in most lizard body surfaces: the dorsal cryptic coloration, the longwavelength based belly and throat, and the ultraviolet (UV) patches found on the outer ventral scales (OVS). Overall, P. liolepis atrata (mainly dorsal and ventral surfaces) are darker than P. liolepis liolepis. This is probably due to a general increase in melanin deposition, a common phenomenon in insular evolution. Also dorsal and ventral coloration are more chromatically pure and short‐wavelength biased in P. liolepis liolepis than in P. liolepis atrata. The colour of throat and belly varies in the same way between populations. Although the UV patches found in OVS are the less variable colour character in this species (probably because they evolve under a strong social selection), we found a highly significant difference in hue, the OVS of male P. liolepis liolepis being more short‐wavelength biased than those of P. liolepis atrata. The available evidence suggests that the hue of UV patches is related to individual quality signalling. Thus, differences in this variable may indicate differences in the intrasexual selection strength.

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Ultraviolet (UV) vision and UV colour patches have been reported in a wide range of taxa and are increasingly appreciated as an integral part of vertebrate visual perception and communication systems. Previous studies with Lacertidae, a lizard family with diverse and complex coloration, have revealed the existence of UV-reflecting patches that may function as social signals. However, confirmation of the signalling role of UV coloration requires demonstrating that the lizards are capable of vision in the UV waveband. Here we use a multidisciplinary approach to characterize the visual sensitivity of a diverse sample of lacertid species. Spectral transmission measurements of the ocular media show that wavelengths down to 300 nm are transmitted in all the species sampled. Four retinal oil droplet types can be identified in the lacertid retina. Two types are pigmented and two are colourless. Fluorescence microscopy reveals that a type of colourless droplet is UV-transmitting and may thus be associated with UV-sensitive cones. DNA sequencing shows that lacertids have a functional SWS1 opsin, very similar at 13 critical sites to that in the presumed ancestral vertebrate (which was UV-sensitive) and other UV-sensitive lizards. Finally, males of Podarcis muralis are capable of discriminating between two views of the same stimulus that differ only in the presence/absence of UV radiance. Taken together, these results provide convergent evidence of UV vision in lacertids, very likely by means of an independent photopigment. Moreover, the presence of four oil droplet types suggests that lacertids have a four-cone colour vision system.

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Many animals display complex colour patterns that comprise several adjacent, often contrasting colour patches. Combining patches of complementary colours increases the overall conspicuousness of the complex pattern, enhancing signal detection. Therefore, selection for conspicuousness may act not only on the design of single colour patches, but also on their combination. Contrasting long- and short-wavelength colour patches are located on the ventral and lateral surfaces of many lacertid lizards. As the combination of long- and short-wavelength-based colours generates local chromatic contrast, we hypothesized that selection may favour the co-occurrence of lateral and ventral contrasting patches, resulting in complex colour patterns that maximize the overall conspicuousness of the signal. To test this hypothesis we performed a comparative phylogenetic study using a categorical colour classification based on spectral data and descriptive information on lacertid coloration collected from the literature. Our results demonstrate that conspicuous ventral (long wavelength-based) and lateral (short wavelength-based) colour patches co-occur throughout the lacertid phylogeny more often than expected by chance, especially in the subfamily Lacertini. These results suggest that selection promotes the evolution of the complex pattern rather than the acquisition of a single conspicuous colour patch, possibly due to the increased conspicuousness caused by the combination of colours with contrasting spectral properties.

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Iberian and North African Podarcis wall lizards constitute a cryptic species complex for which different morphological studies have produced largely conflicting results regarding the number, taxonomic rank and distribution of evolutionary entities. Recently, the assessment of mitochondrial DNA (mtDNA) diversity demonstrated the existence of highly differentiated and geographically consistent clusters, some of which corresponded neatly to partitions previously suggested by morphological analyses. As a result of this concordance, systematic reevaluations were carried out, and at present five different species are recognised: P. bocagei, P. carbonelli, P. vaucheri, P. hispanica and P. atrata, although the latter has been shown to be closely related to one of the various phylogroups that constitute the polytypic (and, from a mitochondrial DNA perspective, paraphyletic) P. hispanica. Given this framework, the first goal of this thesis was to characterize in more detail the dynamics of genetic differentiation among forms of this species complex. Previous descriptions of genetic variability within the complex relied on a single marker – mitochondrial DNA –, and biogeographical inferences were heavily dependent upon poorly supported estimates of relationships. On one hand, we aimed at obtaining a more robust mtDNA-based phylogenetic tree in order to validate previous hypotheses regarding the biogeographical processes governing variability within the clade. On the other hand, we sought to complement these data with information from the nuclear genome, in order to independently assess the distinctiveness of the various clades previously described. Additionally, by examining patterns of nuclear-mitochondrial concordance and analysing in detail the dynamics of a contact zone between two species, we were expecting to evaluate levels of reproductive isolation among forms of the species complex. The reevaluation of mitochondrial phylogenetic relationships was accomplished by selecting a few individuals from each lineage and extending the amount of included sequence data. In addition to the two mtDNA gene regions already published, we sequenced three other fragments, yielding a total of 2425bp. A robust phylogenetic tree was recovered, with a significant proportion of nodes receiving bootstrap support close to 100%. This rigorous assessment of mtDNA relationships between lineages confirmed some of the results obtained in previous studies, such as the paraphyly of P. hispanica. However, other aspects of the phylogeny differ, offering alternative scenarios for the timing and polarity of the colonization of North Africa by wall lizards. In particular, we suggest that the opening of the Strait of Gibraltar might have caused vicariant isolation of Iberian from North African forms, and that a subsequent transmarine colonization could have occurred from North Africa to Iberia. An additional important contribution of this study was the detection of a previously undescribed lineage inhabiting southern Spain, which further exemplifies the evolutionary complexity within this group. Two major lines of research were pursued regarding the description of nuclear gene variation: allozyme electrophoresis and sequencing of nuclear introns. The study of allozyme variation in 30 populations representing all known mtDNA lineages documented a high degree of genetic divergence between most of them, with groups within P. hispanica generally presenting similar levels of differentiation to those observed between fully recognised species. Although this constitutes evidence corroborating the validity of mtDNA lineages as evolutionary distinct units, the application of recent model-based clustering approaches revealed some important discordant patterns with respect to the mtDNA, which may be indicative of the occurrence of gene flow between forms. In this context, two hybrid individuals between the broadly sympatric P. bocagei and P. hispanica type 1 were detected. Summary viii Conversely, the study of two nuclear gene genealogies (β-fibint7 and 6-Pgdint7), demonstrated that, in general, species and mtDNA lineages are strikingly nonmonophyletic, which was unexpected given the high levels of differentiation detected by the use of other markers. An important issue that required clarification was, then, if this pattern emerged as the result of incomplete lineage sorting of ancestral polymorphism (motivated by the four-times higher effective population size of nuclear genes when compared to the mtDNA) or if it resulted from high levels of gene flow. In order to disentangle between these two non- mutually exclusive hypotheses, we estimated levels of gene flow between all pairs of lineages using recent methods of analyses based on coalescent theory that allow the study of the dynamics of admixture and divergence in the same framework. In contrast to the information obtained using classic, F-statistic-based estimators of gene flow, this approach revealed that only a few pairs of lineages have exchanged genes since their divergence, and even fewer show important levels of gene flow, suggesting that in general the various lineages are overall differentiated. Therefore, although gene flow has certainly played an important role in the evolution of the species complex (and as a result some mtDNA lineages may have even lost their nuclear identity), the polyphyletic pattern observed in nuclear genealogies is especially influenced by the persistence of ancestral polymorphism. These studies addressed the subject of gene flow among forms superficially, mostly based on the patterns of mitochondrial/nuclear discordance. In order to characterize in more detail the dynamics of gene flow, a battery of nuclear and mitochondrial markers was studied along a transect crossing the contact zone between P. bocagei and P. carbonelli. This information was complemented with analyses of morphology and fertility. Applying model-based individual multilocus genotype clustering approaches, we document abundance of individuals showing signs of admixture in the locality where the two species meet. However, strong Hardy-Weinberg and linkage disequilibria were observed. This clear bimodality suggests the existence of strong barriers to gene flow, the nature of which still remain obscure. For example, our data suggests neither obvious reductions in hybrid fertility nor the verification of Haldane’s rule. However, divergent natural selection acting on morphology is suggested by the fact that genetically admixed individuals are clearly assigned to one species or the other based on morphological characters. Bimodality within a hybrid zone is usually suggestive of a nearly complete process of speciation; while these results cannot be promptly generalized to other contact zones, they are in accordance with previous suggestions that, although Podarcis lineages are not fully reproductive isolated, levels of gene flow among them do not seem to challenge the ongoing process of differentiation. A second major goal of this thesis was to describe the phylogeographic structure within selected forms of the species complex using multiple molecular markers. In particular, we attempted to evaluate the response of the species to the Pleistocene climatic oscillations, comparing the patterns detected with phylogeographic scenarios described for other species occupying the same regions, and interpreting contrasting signatures left by the Ice Ages on distinct species of Podarcis. The study of mtDNA variation in P. bocagei, P. carbonelli and P. vaucheri, three species with parapatric distributions that replace each other along a latitudinal gradient, allowed us to test two predictions relative to the differential influence of Quaternary climatic oscillations on distinct latitudes: i) northerly distributed species are expected to bear lower levels of genetic subdivision and diversity than species distributed in the south because they were able to survive in fewer, smaller and less stable patches of favourable habitat during glaciations; ii) species distributed in the south should retain signs of long-term effective population size stability, while northern species, because they were more Summary ix confined during glacial stages, should have experienced a rapid demographic growth concomitant with a post-glacial colonization of newly available habitats. Our results show that P. bocagei presents remarkably low levels of diversity and subdivision, a shallow coalescent history (the estimated coalescence time was ~100,000-~70,000 years) and a strong signature of demographic growth. P. vaucheri, on the other hand, presents large levels of genetic diversity even at small geographic scales, strong geographic subdivision, no evidence of demographic growth and an ancient coalescence time, probably dating back to the initial stages of the Pleistocene. The intermediately distributed P. carbonelli presents average values of all studied variables. Taken together, these results entirely fit to our main predictions and demonstrate that different effects of the Ice Ages can be detected at small geographic scales. The analyses of nuclear markers (allozymes and a set of microsatellite loci specifically developed during this study) in populations of P. bocagei and P. carbonelli confirmed, on one hand, the low levels of population subdivision observed in both species from the study of mitochondrial DNA. Moreover, levels of differentiation are higher among populations of P. carbonelli than among populations of P. bocagei, which is also in accordance with higher levels of persistence and a consequent longer time for differentiation in the former inferred from mtDNA. In P. bocagei, we report a progressive loss of genetic diversity in a northwards fashion, consistent with a recent geographic expansion from a reduced source as suspected from mtDNA-based evidence of a rapid demographic growth. Moreover, based on a spatial interpolation of genetic data and on a Bayesian model-based multilocus genotype clustering algorithm, we were able to identify probable expansion routes and to pinpoint with some degree of certainty the area that has probably functioned as a glacial refugium for the species during the last glaciations. Applying the same methodology in P. carbonelli we document some degree of association between genetic clusters and geography, but with only partial correspondence to inferences based on mtDNA; to explain these discordant patterns, we hypothesise that recent fragmentations in the species’ distribution might have led to a loss of the genetic signatures that are typically found in co-distributed species, and that the patterns that are presently observed are more a byproduct of recent genetic drift than of Pleistocene climatic oscillations. Globally, these results constitute a major improvement regarding previous knowledge on evolutionary relationships, stages of differentiation, levels of gene flow and patterns of intraspecific subdivision in Iberian and North African Podarcis. At a more general level, the large-scale study of nuclear gene variation on these lizards, documenting polyphyly and gene flow among mtDNA lineages, illustrates that these correspond to very incipient species with permeable boundaries, highlighting the need for the assessment of multiple genetic markers prior to any taxonomic reevaluations. Moreover, the results have a double importance for the understanding of the Iberian and North African Quaternary biogeography: on one hand, by helping to confirm the transversality of patterns that appear in response to climatic fluctuations, such as glacial fragmentation and post-glacial expansion; on the other hand, by documenting how species-specific such patterns may be even among closely-related species.

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Combining different sources of information is essential for a complete understanding of the process of genetic dif- ferentiation between species. The Iberian and North African wall lizard ( Podarcis ) species complex has been the object of several studies regarding morphological and mitochondrial DNA variation but, so far, no large-scale survey of nuclear variation within this group has been accomplished. In this study, ten polymorphic allozyme loci were stud- ied in 569 individuals collected across the Iberian Peninsula and North Africa. The obtained data were analysed using both conventional population genetic tools and recent Bayesian model-based clustering methods. Our results show that there are several well-differentiated entities corroborating the major splits observed in mtDNA analyses. These groups correspond not only to the fully recognized species Podarcis bocagei , Podarcis carbonelli , and Podarcis vaucheri but also to multiple forms within the polytypic Podarcis hispanica , all of which have a similar level of dif- ferentiation to that observed between the acknowledged species. However, relationships between forms are weakly supported both by population and individual clustering methods, suggesting a scenario of a rapid diversification that contrasts to the clear bifurcating model assumed from previous mtDNA analyses. Individual multilocus analyses report few individuals misassigned or apparently admixed, some of which are most likely explained by the persis- tence of high levels of ancestral polymorphism. Other admixed individuals, however, are probably the result of lim- ited levels of gene flow between forms.

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Background: The study of recently-diverged species offers significant challenges both in the definition of evolutionary entities and in the estimation of gene flow among them. Iberian and North African wall lizards (Podarcis) constitute a cryptic species complex for which previous assessments of mitochondrial DNA (mtDNA) and allozyme variation are concordant in describing the existence of several highly differentiated evolutionary units. However, these studies report important differences suggesting the occurrence of gene flow among forms. Here we study sequence variation in two nuclear introns, β-fibint7 and 6-Pgdint7, to further investigate overall evolutionary dynamics and test hypotheses related to species delimitation within this complex. Results: Both nuclear gene genealogies fail to define species as monophyletic. To discriminate between the effects of incomplete lineage sorting and gene flow in setting this pattern, we estimated migration rates among species using both FST-based estimators of gene flow, which assume migration-drift equilibrium, and a coalescent approach based on a model of divergence with gene flow. Equilibrium estimates of gene flow suggest widespread introgression between species, but coalescent estimates describe virtually zero admixture between most (but not all) species pairs. This suggests that although gene flow among forms may have occurred the main cause for species polyphyly is incomplete lineage sorting, implying that most forms have been isolated since their divergence. This observation is therefore in accordance with previous reports of strong differentiation based on mtDNA and allozyme data. Conclusion: These results corroborate most forms of Iberian and North African Podarcis as differentiated, although incipient, species, supporting a gradual view of speciation, according to which species may persist as distinct despite some permeability to genetic exchange and without having clearly definable genetic boundaries. Additionally, this study constitutes a warning against the misuse of equilibrium estimates of migration among recently-diverged groups.

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Automated image classification is a thriving field of machine learning, and various successful applications dealing with biological images have recently emerged. In this work, we address the ability of these methods to identify species that are difficult to tell apart by humans due to their morphological similarity. We focus on distinguishing species of wall lizards, namely those belonging to the Podarcis hispanicus species complex, which constitutes a well-known example of cryptic morphological variation. We address two classification experiments: (1) assignment of images of the morphologically relatively distinct P. bocagei and P. lusitanicus; and (2) distinction between the overall more cryptic nine taxa that compose this complex. We used four datasets (two image perspectives and individuals of the two sexes) and three deep-learning models to address each problem. Our results suggest a high ability of the models to identify the correct species, especially when combining predictions from different perspectives and models (accuracy of 95.9% and 97.1% for females and males, respectively, in the two-class case; and of 91.2% to 93.5% for females and males, respectively, in the nine-class case). Overall, these results establish deep-learning models as an important tool for field identification and monitoring of cryptic species complexes, alleviating the burden of expert or genetic identification.

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Les Reptiles des Pyrénées traite en détail les 32 espèces protégées présentes sur l’ensemble de la chaîne (France, Espagne et Andorre). Cette faune herpétologique est le résultat d’une entreprise natu­raliste combinant enquête bibliographique et reportage photographique. Elle propose une vaste synthèse de données relatives à plusieurs champs disciplinaires – systématique, ­taxi­nomie, biogéographie, écologie, biologie… – jusque-là dispersées dans plusieurs centaines d’articles et ouvrages. Les variations phénotypiques et les particularités écologiques des serpents, lézards et tortues occupant l’espace pyrénéen y sont illustrées par de très nombreuses photo­graphies exclusivement réalisées in situ, de 2 m à 3 143 m d’altitude. Des cartes de répartition précises (mailles UTM 10 km × 10 km), basées sur des sources scientifiques, complètent et éclairent le propos biogéographique. Les espèces et sous-espèces endémiques ou subendémiques de la chaîne, de même que celles qui y ont un statut particulier (très localisées, vulnérables…), ont fait l’objet d’une attention par­ti­cu­lière et les menaces qui pèsent sur les reptiles des Pyrénées sont largement exposées. Cet ouvrage sera précieux pour les pyrénéistes, naturalistes et gestionnaires d’espaces naturels de la chaîne (parc national, réserves, sites Natura 2000, parcs naturels régionaux…), qui disposeront là d’une mine d’informations sur le sujet.

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Comparisons among eight populations of Podarcis lilfordi, seven of Podarcis pityusensis (representative lizards of the Balearic Archipielago), and one of Podarcis hispanica atrata fro the Columbretes Islands, have been made from a morphological point of view, measuring 15 morphometric characters of the lizards. Cluster and discriminant analyses were obtained for males and females to show the relationships among the different populations and species. These analyses gave three groups: one of them with the most P. lilfordi populations, other with the most P. pityusensis populations, and a well seperated group with the P. hispanica atrata species. Some populations connect the groups of P. lilfordi and P. pityusensis. The possibility that P. lilfordi and P. pityusensis be two pattern of variation inside the same species instead of two well differentiated species is considered, altrough further studies are necessary.

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Cytoplasmic markers are often used alone to reconstruct phylogenetic and phylogeographic patterns. Evolutionary, demographic and ecological processes that cause these patterns are most often investigated using nuclear markers. However, some mechanisms, like introgressive hybridization, may result in gene histories differing from species histories. Further, genetic consequences of introgressive hybridization may be particularly marked for cytoplasmic markers comparatively to nuclear markers. This is why the reliability of cytoplasmic markers as a tool for pattern reconstruction is currently debated. To evaluate the value of cytoplasmic markers for pattern reconstructions and process studies, we compared, within the species complex of Iberian Wall Lizards Podarcis hispanicus, the evolutionary history inferred from a mitochondrial marker with the history inferred from nuclear markers, and in fig species of section Galoglychia, the history inferred from chloroplast markers with the history inferred from nuclear markers. In both cases, we evidenced several cases of cyto-nuclear discordances caused by ancient events of introgressive hybridization. In the lizard example, discordance was caused by the complete replacement of the mitochondrial lineage of one species, in a large part of its distribution range, by a mitochondrial lineage belonging to another species currently extinct or not described yet. In the Ficus example, congruence between plant chloroplast phylogeny and pollinator phylogeny supports the hypothesis that host shifts are responsible of both cyto-nuclear discordance in plant phylogenies and pollinator-plant phylogenetic incongruence. Both cases demonstrate that, because of introgressive hybridization, cytoplasmic markers are of limited interest to precisely describe phylogenetic and phylogeographic patterns. However, these introgressive hybridization events make cytoplasmic markers useful to investigate processes.

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Mitochondrial markers are still often used alone to identify evolutionary units, despite widespread evidence for processes such as incomplete lineage sorting or introgressive hybridization that may blur past population history. The combination of mitochondrial DNA data with other sources of information (morphology, nuclear genes) is a powerful tool to reveal when and why mitochondrial markers are potentially misleading. In this study, we evaluate the performance of mtDNA markers to unravel the evolutionary history of Spanish lizards from the Podarcis hispanicus species complex. We first uncover several cases of discordance between morphological and mitochondrial data in delimitation of taxa. To assess the origin of these discordances, we analysed the same populations using several independent nuclear loci. Both morphological and nuclear markers identified the same three evolutionary units in the region, while mitochondrial data revealed four deeply divergent lineages. We suggest here that the most likely scenario to explain this discordance is ancient mitochondrial introgression originating from a fourth evolutionary unit presently absent from the study area. Notably, this resulted in a complete replacement of the original lineage in a large part of the distribution of one of the taxa investigated. We discuss the potential evolutionary scenarios leading to this complete mitochondrial replacement and suggest why the previous studies have failed to recover the correct history of this species complex.

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Recent genetic works have suggested that the Iberian wall lizard Podarcis hispanicus (Steindachner, 1870) sensu lato is in fact a species complex. Several forms have already been elevated to species rank, including the north-eastern Spanish form alternatively named Podarcis atratus (Boscá, 1916) or P. liolepis (Boulenger, 1905) in different works. However, a recent study has demonstrated that the province of Valencia, where most individuals of the type series of Lacerta muralisvar. liolepis Boulenger, 1905 originate from, is inhabited by populations that are conspecific with the north-eastern Spanish form. Consequently the nomen liolepis has precedence over atratus to designate the north-eastern species of the P. hispanicus complex which should thus bear the name Podarcis liolepis.

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A total of 35 morphological characters (biometry, scalation, chromatic pattern) were studied through multivariate analyses on 10 populations sampled across the range of the Iberian wall lizard (Podarcis hispanica) in Portugal. Biometry clearly splits the samples into two different types. Differences in scalation between the two types were not clear, but multiple correspondence analyses showed that different chromatic patterns Ž t each of the types: one presented dark dorsal patterns (e.g. reticulated, eyeled, striped) and whitish-pearly belly, while the other showed green or yellow-brown patterns and yellow-orange belly. These two morphotypes constitute different molecular lineages and have different ranges of distribution.

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Changes in habitat structure constitute a major factor explaining responses of reptiles to fire. However, few studies have examined habitat factors that covary with fire-history variables to explain reptile responses. We hypothesise that more complex habitats should support richer reptile communities, and that species-specific relative abundance should be related to particular habitat features. From spring 2012–2014, twenty-five transects were surveyed in the Albera Region (north-east Iberia). The vegetation structure was measured and the extent of habitat types in a 1000-m buffer around each transect calculated. Reptile-community metrics (species richness and reptile abundance) were related to fire history, vegetation structure, and habitat types, using generalized additive models. These metrics correlated with habitat-structure variables but not with fire history. The number of species increased with more complex habitats but decreased with pine-plantation abundance in the 1000-m buffer. We found contrasting responses among reptiles in terms of time since fire and those responses differed according to vegetation variables and habitat types. An unplanned fire in August 2012 provided the opportunity to compare reptile abundance values between pre-fire and the short term (1–2 years) after the fire. Most species exhibited a negative short-term response to the 2012 fire except Tarentola mauritanica, a gecko that inhabits large rocks, as opposed to other ground-dwelling species. In the reptiles studied, contrasting responses to time since fire are consistent with the habitat–accommodation model of succession. These differences are linked to specific microhabitat preferences and suggest that functional traits can be used to predict speciesspecific responses to fire.

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Under the so-called PEIN (Natural Interest Sites Plan, declared by the catalan government) a in the south of the of Lleida demarcation (NE Iberian Peninsula) the herpetological species observed in recent years are shown. There are a total of 8 protected areas sampled, all belonging to the semi-arid continental scrubland region, below the first slopes of the pre-Pyrenees. In one of these areas there is a wetland with a reservoir. For all the areas sampled the data of localization, in UTM 10x10 sq. Km, and date of observation of the different species is given. A total of 23 species have been found in the different habitats visited.

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In addition to species of the North Spanish herpetofauna the allochthonous and wellestablished populations of the Ibiza wall lizard Podarcis pityusensis and the Italian wall lizard Podarcis siculus were observed at Spain`s Atlantic coast.

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Andorra was visited two times in August. A survey of landscape, vegetation, and climate of Andorra is given. Six species of lacertid lizards (Lacerta agilis garzoni, Podarcis liolepis, Podarcis muralis brongniardii, Psammodromus algirus, Timon lepidus lepidus, Zootoca vivipata louislantzi) could be found. Mainly information on distribution and habitat observations are reported.

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SUMMARY Increasing human population, international commerce and mobility are accelerating the introduction of biota to a formerly unknown extent. Consequently, intended and unintended introductions of non-native species are recognized as a severe problem in nature conservation. The Common wall lizard (Podarcis muralis), a species with a sub-Mediterranean native distribution, has established more than 150 non-native populations in Central Europe. This species represents an ideal model system to examine the role of the source region on establishment success, to assess the genetic consequences of biological invasions, and to study the consequences of intraspecific hybridization on native wall lizard lineages. I performed a ‘top-down approach’, aiming to identify the prerequisites first, and subsequently narrowing the perspective towards potential causal mechanisms and evolutionary processes. The adaptive capabilities and wide distribution of many species implicate that the quality and breadth of ecological niches vary across time, space and therefore among populations and evolutionary lineages. To test this hypothesis, I analysed for the first time whether climatic niches differ at the intraspecific level in invasive species (Chapter I). I assigned 77 introduced populations in Central Europe to eight geographically distinct evolutionary lineages using DNA barcoding. The resulting dataset was used in combination with species distribution models (SDMs) based on climatic information from native and invasive ranges to test for intraspecific niche divergence. The analysed lineages had rather similar realised niches in their native and invasive ranges, whereas inter-lineage niche differentiation was comparatively strong. However, I found only a weak correlation between geographic origin (i.e. mtDNA-lineage) and invasive occurrence. Lineages with narrow realised niches still became successful invaders far outside their native range, most probably due to broader fundamental niches. The results indicate that the observed niche differentiation among evolutionary lineages is mainly driven by niche realisation and not by differences in the fundamental niches. Furthermore, the findings suggest that cryptic niche conservatism may in general explain the invasion success of species in areas with apparently unsuitable climate. The wide applicability of genetic tools in conservation biology requires the development of minimalinvasive or non-invasive DNA sampling techniques. To adequately address questions of population genetics in this thesis ca. 800 individuals had to be genotyped. Since tail-clipping harms the locomotory performance and energy reserves of wall lizards, I compared the quantity and quality of the isolated DNA collected from buccal swabs and tail tips in Chapter II. These results show that buccal swabs are a simple and efficient non-invasive sampling method for DNA analysis in adult lacertid lizards. In Chapter III, I determined the geographic origin of the northernmost introduced population of the Italian wall lizard Podarcis siculus in Rapperswill, Switzerland. This population originated from a region between the Po plain (Italy) and the northern Adriatic region (Croatia). Chapter IV comprises the first record of P. liolepis introduced in Germany within a syntopic population of its congener P. muralis. The founder individuals presumably stem from a region in the eastern Pyrenees, where sympatric populations are known. In concordance with behavioural observations, no evidence for gene flow between the two species was found. Compared to the analysed native populations, high levels of genetic diversity have been retained in the introduced population of both species. Erklärungen 148 Chapter V provides the first evidence for intraspecific hybridisation between native and introduced wall lizard lineages along the Upper Rhine Rift. As a devastating consequence of introductions, a rapid genetic assimilation of native wall lizard populations caused by strong introgression from introduced Italian lineages could be demonstrated. The genetic diversity of hybrid populations was substantially higher than in introduced and native populations belonging to a single lineage but, in contrast to previous studies, the relationship between genetic diversity and admixture level was non-linear and rapidly reached a plateau of high genetic diversity at an admixture level of two. However, even introduced populations with low founder sizes and from one source population retained moderate levels of genetic diversity and no evidence for a genetic bottleneck was found. The extent of introgression and the dominance of alien haplotypes in mixed populations indicate that introductions of non-native lineages represent a significant threat for the genetic integrity of native populations due to the rapid creation of hybrid swarms. Fine scale genetic analyses of evolutionary processes acting at the invasion front of expanding populations have rarely been performed at a local scale. I analysed spatial patterns of genetic variation within an expanding introduced wall lizard population in Passau (Chapter VI). The results demonstrate that significant genetic population structure can emerge rapidly at a small spatial scale. The genetic differentiation tended to increase, while the genetic diversity declined from the centre of introduction towards the expanding range margin. I assume that the pronounced territoriality of P. muralis, which results in ’high rates‘ of noncontiguous and stratified dispersal from longer established sites, is sufficient to maintain genetic diversity despite founder events at the invasion front. The final Chapter VII summarises most of the important findings of this thesis in order to provide information for field herpetologists and conservationists. Furthermore, I address the difficult phenotypic assignment of populations to evolutionary lineages and discuss the problem of how to deal with invasive populations in the light of the current conservation legislation.

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The Common Wall Lizard (Podarcis muralis) has been introduced within large parts of Central Europe, the UK and parts of North America. In an introduced population of this species in Lower Saxony, Germany, we found in addition to mtDNA haplotypes of P. muralis also haplotypes of its congener Podarcis liolepis, a species that hitherto has never been recorded outside its native range. We therefore, (1) wanted to identify the geographic origin of the founder individuals of both non-native populations, (2) test for hybridization between introduced individuals of both species in Germany and (3) compare levels of genetic diversity between native and introduced populations.We sequenced a fragment of the mitochondrial cytochrome b gene and genotyped individuals of the introduced as well as native populations of both species at eleven microsatellite loci. Our results suggest that the founders presumably stem from a region in the eastern Pyrenees, where sympatric populations of P. muralis and P. liolepis are known. No evidence for gene flow between the two species was found in the introduced population. These results are consistent with behavioural observations indicating agonistic interactions of P. muralis towards P. liolepis rather than cross-species attraction. Compared to the native populations, high levels of genetic diversity have been retained in the introduced population of both species and no evidence for a genetic bottleneck was found. The effective population size was high in P. muralis, but substantially smaller in P. liolepis.

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In recent years, great attention has been paid to many Podarcis species for which the observed intra-specific variability often revealed species complexes still characterized by an unresolved relationship. When compared to other species, P. siculus underwent fewer revisions and the number of species hidden within this taxon may have been, therefore, underestimated. However, recent studies based on genetic and morphological data highlighted a marked differentiation of the populations inhabiting the western Pontine Archipelago. In the present work we used published genetic data (three mitochondrial and three nuclear gene fragments) from 25 Podarcis species to provide a multilocus phylogeny of the genus in order to understand the degree of differentiation of the western Pontine populations. In addition, we analyzed new morphometric traits (scale counts) of 151 specimens from the main islands of the Pontine Archipelago. The phylogenetic analysis revealed five principal Podarcis groups with biogeographic consistency. The genetic distinctiveness of the Podarcis populations of the western Pontine Islands is similar or even more ancient than those observed in numerous other pairs of Podarcis sister species. In the light of these evidences we raise the western Pontine lizards to specific rank; thus they should be referred to as Podarcis latastei.

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Whereas the range size of endangered species is undoubtfuly useful to predict risk of extinction, the role of their life-history characteristics is much less clear, and their effects may depend on the nature of the threatening factors. Such factors, for instance, are known to be different on islands and on the mainland. We used phylogenetically based statistical analyses to study the relationships among conservation status, insularity, range size, and life-history traits in a clade of Western Palaearctic lacertids including insular and continental species. These lizards are ecologically similar, but they show wide variation in life-history traits and vulnerability to extinction. Insular species of a given size had smaller clutches than mainland ones. Degree of threat was best predicted by a logistic regression including range size, insularity, clutch size, and the insularity × clutch size interaction. On the mainland, but not on islands, threatened species had smaller clutches than non-threatened ones. On islands, small clutch size is probably an adaptive trait, and it might predispose certain species to extinction, but the intrinsic characteristics of such species remain unclear. However, small clutch size was a good predictor of extinction risk on the mainland, having evolved most frequently in late maturing species from montane habitats in which climatic conditions limit their reproductive output and increase their vulnerability to stochastic hazards or habitat fragmentation.

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Aim: The Mediterranean basin has a long history of interactions between humans and biota, with multiple ancient and recent introductions of alien species. Such a multitude of introductions makes it difficult to distinguish between alien and native species but provides an excellent opportunity to investigate factors related to intro ‐ductions and long‐term persistence of alien species. In this study, we combined ge ‐netic and distribution data to identify the factors promoting the presence of alien reptiles on islands, considering human‐related, geographic and species features. Furthermore, we assessed whether the use of genetic evidence to identify alien spe ‐cies improves inference of the factors determining their distribution.Location: Mediterranean islands.Methods: We combined genetic data and distribution databases to obtain informa‐tion on biological traits and on the native/alien status of reptiles on >900 Mediterranean islands, and we gathered data on geographic and human features of islands. We then used spatially explicit generalized additive mixed models to identify the factors associated with the establishment of alien reptiles.Results: Alien reptile populations are more frequent on islands far from the native range and with large human population. Alien populations of reptiles that are able to feed on plants are particularly frequent. Traditional data sources underestimate the frequency of alien reptiles on Mediterranean islands, and using genetic evidence to assess the status of populations provided a more complete picture of the factors as ‐sociated with the presence of alien populations.Main conclusions: Humans are key drivers of the distribution of alien reptiles on Mediterranean islands, but the distributions are determined by a complex interplay between human activities, geographic factors and species features. Genetic data are essential for obtaining reliable biogeographic assessments of invasive species, par ‐ticularly in systems with a long history of human influence.

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Research on the taxonomy of European amphibians and reptiles has increased noticeably over the last few decades, indicating the need for recognition of new species and the cancellation of others. This paper provides a critical review of recent changes and draws up a tentative species list.

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The last species list of the European herpetofauna was published by Speybroeck, Beukema and Crochet (2010). In the meantime, ongoing research led to numerous taxonomic changes, including the discovery of new species-level lineages as well as reclassifications at genus level, requiring significant changes to this list. As of 2019, a new Taxonomic Committee was established as an official entity within the European Herpetological Society, Societas Europaea Herpetologica (SEH). Twelve members from nine European countries reviewed, discussed and voted on recent taxonomic research on a case-by-case basis. Accepted changes led to critical compilation of a new species list, which is hereby presented and discussed. According to our list, 301 species (95 amphibians, 15 chelonians, including six species of sea turtles, and 191 squamates) occur within our expanded geographical definition of Europe. The list includes 14 non-native species (three amphibians, one chelonian, and ten squamates).

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The naming of species and of all systematic entities of living things are dynamic con- cepts. Through scientific research, changes in the systematics and names of amphibi- ans and reptiles are constantly being pro- posed, much to the chagrin of many profes- sional and amateur herpetologists. Yet most changes are necessary if taxonomy and systematics are to reflect evolutionary his- tory and phylogeny, rather than letting user–friendliness and conservatism prevail. The European herpetofauna and its taxo- nomy have received increasing attention through molecular (e.g. DNA) studies. Once most animal groups have been studied in this way, stability may be established. In the following pages, a concise overview of changes in the taxonomy of European amphibians and reptiles is presented. We have restricted ourselves to the geographi- cal boundaries of Europe, albeit excluding former Soviet states. Thus, politically Euro- pean areas in Asia (e.g. Cyprus and Greek islands in front of the Aegean Turkish coast) and Africa (e.g. Canary Islands and Ma- deira) have not been included. For a time frame we have used the species and names as presented in the well–known European field guide by ARNOLD (2002), although numerous changes were proposed and published prior to 2002. We refer to the cited literature for more comprehensive explanations. Only species or higher level changes concerning European taxa are listed. Subspecific changes and intraspeci- fic variability are noted only when contra- dicting long–established monotypy of a species, or when subspecies are being rejected. Vernacular names are mostly adopted from ARNOLD (2002). Some exo- genous species that are well–established and are reproducing on European soil, are included and are listed separately. Species believed to have been introduced to Europe over 100 years ago and persisting until today, have been included in the list of endogenous species. Many other non– native species have, however, been en- countered in the wild in Europe. Final content changes were made on De- cember 1, 2007. This type of update and the proposed species list will most likely be outdated at its date of publication. Further updating by means of a consecutive series of similar papers seems desirable.

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From 2012 to 2021, a total of 435 observations of reptiles were collected in the Special Area of Conservation of the Bardenas Reales (Navarre) and its surrounding area, corresponding to 15 autochthonous species. The observations were mapped on 5 x 5 km UTM grids, which allowed the distribution of these species in the area to be updated. Their pattern of presence can be summarised in three types of spatial occupation: ubiquitous species, species with a specific distribution in selective environments and species concentrated in restricted and specific areas. The Mediterranean scrub is the habitat with the highest number of observations (63 % of the records). The most widely represented lizard species was Psammodromus algirus (Linnaeus, 1758) (73.81 % of the squares), and the snake species, Natrix maura (Linnaeus, 1758) (38.10 %). Emys orbicularis (Linnaeus, 1758), favoured by a population monitoring and habitat improvement programme, was located in 16.67 % of the squares. The rarest species were Chalcides bedriagai (Boscá, 1880) (2.38 %) and Vipera latastei Boscá, 1878 (9. 52 %).

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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 Podarcis 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 frequently 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.

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Locomotor behaviour varies between two subspecies of the Spanish wall lizard Podarcis hispanica. One subspecies inhabits the Columbretes islands, the other lives on the Spanish mainland. Size standardized voluntary speeds (as measured in unrestrained laboratory conditions) are lower in the island population (P. h. atrata) than in the mainland population (P. h. hispanica). Maximal running performance (when chased) is much higher in the mainland population than in the island population. High speed video recordings show that subspecies differ in gait characteristics: individuals from the mainland modulate running velocity primarily by modifying stride length, individuals from the island primarily by altering stride frequency. P. h. hispanica’s strategy for modulating speed probably allows this mainland subspecies to attain higher maximal speeds than the island subspecies P. h. atrata. Theoretical considerations suggest that at high speeds, P. h. hispanica’s running style is energetically more favourable, but this hypothesis awaits experimental verification. We suggest that the differencesin locomotion efficiency between the subspecies result from differences in predation pressure between the mainland and the island. The mainland study site has a higher predator diversity and offers less hiding opportunities to the lizards.

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We compared the behavior of two subspecies ofPodarcis hispanica lizards in cages that had been chemically marked by a saurophagous snake, the viperVipera latastei. One of the subspecies (P.h. atrata) has experienced a relaxation from predation pressure by this viper, as snakes were eradicated from the island it inhabits over 100 years ago. Nevertheless, individuals fromP.h. atrata responded to the snake`s chemicals similarly to individuals from a population ofP.h. hispanica, currently sympatric withV. latastei. Lizards exhibited more stress-indicating behaviors (foot shakes, tail vibrations, sudden starts), became less mobile, and tongue-flicked more while moving in a snake-inhabited terrarium than when in a clean, unfamiliar terrarium.

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We tested the hypothesis that an evolutionary trade-off exists between the capacity to run on level terrain and the ability to climb inclined structures in lacertid lizards. Biomechanical and physiological models of lizard locomotor performance suggest that the morphological design requirements of a ground-dwelling vs. scansorial life style are difficult to reconcile. This conflict is thought to preclude simultaneous evolution of maximal locomotor performance on level and inclined terrain. This notion has been corroborated by comparative studies on lizard species from other groups (Anolis, Chamaeleo, Sceloporus), but is not supported by our data on 13 species from the family Lacertidae. We found no indication of a negative association between maximal sprint speed of lizards over a level racetrack (indicative of ground-dwelling locomotor performance), on an inclined stony surface (indicative of climbing performance over rock faces) and inclined mesh surface (indicative of clambering performance among vegetation). Moreover, morphological characteristics associated with fast sprinting capacities (e.g. long hind limbs) apparently enhance, rather than hinder climbing and clambering performance. We conclude that in our sample of lacertid lizards, the evolution of fast sprinting capacity on level terrain has not inflicted major restrictions on climbing and clambering performance.

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Preferred body temperatures (Tp) were analysed in the lacertid Podarcis vaucheri, from S. Iberia and N. Africa which has been recently revalidated as a full species within the P. hispanica complex. Adult lizards from a Moroccan population were exposed to a thermal gradient and Tp was measured at ten time intervals. Intraspecific Tp variation due to sex and reproductive condition (Tp: non-pregnant females

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During two journeys from 27.04. – 07.05.2008 and from 21.04. – 27.04.2012, I visited several parts pf Spain together with some friends of mine. We focused on the lizards of the Iberiasn Peninsula. Specifically, we wanted to make some beautiful pictures of the Ocellated Lizatrds Timon lepidus and Timon nevadensis, which unfortunately did not always succeed on the first trip. In the following article, I briefly present the lizards of the family Lacertidae that were found during the two journeys together with some of their localities.