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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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Lizard biology is strongly linked to the spatial structure of their environment, since lizards are sedentary and depend on habitat complexity for thermoregulation, hydroregulation and antipredator defence. Agricultural stone walls are their most commonly occupied habitats, providing them with food, shelter and thermoregulating surfaces. Management may affect microhabitat structure, with uniformized concrete walls and the trimming of surrounding vegetation leading to a reduction of the available shelter. Here we analyzed the daily activity patterns and microhabitat use of Podarcis bocagei wall lizards occupying agricultural walls. From May to July 2022, lizard surveys based on visual encounters were performed in corn (Zea mays) fields in Vairão, NW Portugal, following line transects along stone walls. Each observation included information on the lizard class (adult male, female or immature), time of day, height above ground, epiphytic vegetation and number of holes. Associations between the environmental variables were examined through a multiple correspondence analysis. Results show a tendency for different patterns of activity between the morning and afternoon periods, suggesting that lizards thermoregulate during the first hours of the day, avoid activity during the heat of midday, and forage in the afternoon. Immature individuals occupied more open sections of the wall, with small or no holes and sparce vegetation. Males and females show no difference in activity or preferred microhabitat structure. Lizards avoided sections with no vegetation, and were seldom observed at ground level. They were most often observed near smaller holes than larger holes, possibly avoiding these locations due to predation from larger predators such as Timon lepidus. Studies in nonpristine areas suggest guidelines on how reptiles may cope with humanized landscapes and, ultimately, provide clues on how agriculture practices and biodiversity conservation could be better harmonized.

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As part of a wider study examining the impacts of corn pesticides on lacertid lizards in north-western Portugal, we examined various physiological, biochemical, and histological biomarkers of exposure and effect among field populations of Podarcis bocagei. Biomarkers included body condition index, standard metabolic rate, locomotor performance, parasitization, glutathione oxidative pathways and related enzyme activity, lipid peroxidation and liver and testis histology. Few of the various biomarkers investigated provided statistically significant evidence of toxic effect. However, using a weight of evidence approach, we conclude that pesticides are affecting lizards living in the vicinity of pesticide exposed corn agriculture sites. Lizards from these locations present a profile of animals under metabolic stress with reduced condition indices, increased standard metabolic rate, lower incidence of hepatocyte vacuolation, altered iron metabolism, increased activation of GSH oxidation pathways, and even increased prevalence of hemoparasites.

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Mesocosms (i.e., outdoor, man-made representations of natural ecosystems) have seldom been used to study the impact of contaminants on terrestrial ecosystems. However, mesocosms can be a useful tool to provide a link between field and laboratory studies. We exposed juvenile lacertid lizards for a period of over one year to pesticides (herbicides and insecticides) in mesocosm enclosures with the intention of validating field observations obtained in a previous study that examined the effects of corn pesticides in Podarcis bocagei. Our treatments replicated field conditions and consisted of a control, an herbicides only treatment (alachlor, terbuthylazine, mesotrione and glyphosate) and an herbicides and insecticide treatment (including chlorpyrifos). We used a multi-biomarker approach that examined parameters at an individual and sub-individual level, including growth, locomotor performance, standard metabolic rate, biomarkers of oxidative stress, esterases and liver histopathologies. Although mortality over the course of the exposures was high (over 60%), surviving individuals prospered relatively well in the mesocosms and displayed a broad range of natural behaviours. The low numbers of replicate animals compromised many of the statistical comparisons, but in general, surviving lizards exposed to pesticides in mesocosm enclosures for over one year, thrived, and displayed few effects of pesticide exposure. Despite the difficulties, this work acts as an important stepping-stone for future ecotoxicology studies using lizards.

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Lizards are one of the least studied groups in ecotoxicology and despite a recent increase in number of studies, there is still a lack of knowledge regarding their response to environmental contamination. Lacertid lizards have been identified as potential model species for reptile ecotoxicology in Europe. They offer several advantages over other groups as they are usually highly abundant; have strong site fidelity; are easy to capture, mark and recapture in the field; and are easy to maintain in captivity. The main goal of our project is to assess if highly abundant lacertid lizards belonging to the genus Podarcis in Southern Europe, can be used as bioindicators of pesticide exposure and toxicity in areas of intensive agriculture. We have chosen Podarcis bocagei because is a widely abundant species and the only lacertid present on high pesticide usage corn fields dominating the landscape of north- western Portugal. As a first approach, we will document the abundance, demography and biological fitness of the species occurring in areas of either intensive or negligible pesticide usage. A second approach aims to study naïve lizards in more controlled experiments, both in mesocosm and laboratory experiments to validate field observations. We intend to employ a variety of techniques to assess pesticide exposure and toxicity including biochemical, behavioural, morphological and molecular biomarkers. On a larger scale we hope to study bioaccumulation processes in lizards, reflect on the biological significance of the reported tissue concentrations and develop criteria to predict risk to lizard populations. In the future, similar projects using lacertids as bioindicators could be developed in other parts of the Mediterranean Basin.

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At the European level, lacertid lizards have been proposed as potential model species for reptile ecotoxicology. We studied demographic and morphological aspects of natural field subpopulations of Podarcis bocagei inhabiting similar agricultural habitats which were either regularly exposed to pesticides, or not. Parameters examined in this study included population size and density, sex ratio, adult body size, fluctuating asymmetry in femoral pores and parasite prevalence. In general, we detected few statistically significant differences between the exposed and reference subpopulations. Although field situations are ecologically complex and factors other than pesticides may be acting, the absence of observable effects on field subpopulations is probably indicative that lizards are coping or compensating for this level of exposure.

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In Europe, reptiles have been recently included in environmental risk-assessment processes for registration of plant-protection products. However, data on toxicity effects of most compounds are lacking. Chlorpyrifos is the most commonly used organophosphorus insecticide worldwide. In the present study, the authors exposed a lacertid lizard, Podarcis bocagei, to sublethal concentrations of chlorpyrifos. Individuals were exposed through spiked food for a period of 20 d (low dose 0.12 mg/kg/d, high dose 1.57 mg/kg/d). After exposure, various biomarkers of exposure and effect were evaluated, including the activities of glutathione S-transferase and enzymes involved in the glutathione redox cycle, glutathione concentrations, activities of esterases, liver and testes histopathologies, as well as locomotory and predatory behavior. The results indicate that sublethal, subchronic exposure to chlorpyrifos can affect P. bocagei in a dose-dependent manner. Adverse effects occurred at both the subindividual and individual levels, including inhibition of carboxylesterases and cholinesterases (ChEs), liver histopathological changes, and altered predatory behaviors. Animals exposed to chlorpyrifos took more time to capture and subdue prey items. The results suggest a link between effects at subindividual levels of organization with those observed at the whole individual level after exposure to environmentally realistic dosages of chlorpyrifos.

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Lizards are among the least studied groups in ecotoxicology and despite a recent increase in the number of studies, there is still a lack of knowledge regarding their response to environmental contamination. In Europe, lacertid lizards have been identified as potential model species for reptile ecotoxicology. The main question of this project was to assess if a highly abundant lacertid lizard belonging to the genus Podarcis, could be used as a bioindicator of pesticide exposure and toxicity in agricultural areas. To achieve this end, we used a three-stage tiered approach. The first tier took the form of a field survey to document the nature of the exposure and the population parameters of lacertids occurring in areas of intensive pesticide usage as well as areas of negligible pesticide usage. The second tier consisted of a mesocosm study in which naïve lizards were exposed to pesticides in a controlled experiment. Finally, the third tier included a laboratory approach to the effects of one of the most common insecticides used worldwide, chlorpyrifos. We assessed pesticide impact in individuals from the different tiers using a comprehensive set of biomarkers applied at different levels of biological organization, including population parameters as well as behavioral, physiological, biochemical and histological biomarkers. We detected few statistically significant differences between reference and exposed populations in the field study. Confirming the difficulty to isolate the effects of contaminants on natural populations where other local factors, natural cycles or stochastic events occur. P. bocagei populations seem to be able to compensate with the observed level of pesticide exposure. Nevertheless, individuals living in exposed sites seem to be less ecologically fit than those living in reference sites, presenting a depleted nutritional status and signs of metabolic stress. The results from the mesocosm setting reinforce these results. Lizards prospered relatively well in all enclosures, treated or not with pesticides, presenting a broad range of natural behaviors. The laboratory approach confirmed P. bocagei as valuable indicator of sub-lethal exposure to environmental realistic doses of chlorpyrifos. According to our results, P. bocagei seems to be a suitable bioindicator of pesticide exposure.

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Lizards are among the least studied groups in ecotoxicology, and despite a recent increase in the number of studies, there is still a lack of knowledge regarding their response to environmental contamination. In Europe, lacertid lizards have been identiied as potential model species for reptile ecotoxicology. The main question of our project was to assess if highly abundant lacertid lizards belonging to the genus Podarcis, could be used as bioindicator of pesticide exposure and toxicity in agricultural areas. To achieve this end, we used a three- stage tiered approach. The irst tier took the form of a ield survey to document both exposure and population endpoints of lacertids occurring in areas of intensive pesticide and usage and areas of negligible pesticide usage. The second tier was a mesocosm study in which naïve lizards were exposed to pesticides in a controlled experiment. Finally, the third tier included a laboratorial approach to the efects of one of the most common insecticides used worldwide, chlorpyrifos. We assessed pesticide impact using a comprehensive set of endpoints applied at diferent levels of biological organization, including behavioral, physiological, biochemical and histological biomarkers. The ield work conirmed the di culty of diferentiation between the efect of contaminants and other (local) factors at the population level but our results suggest a diference in the metabolic activity between animals from reference and exposed locations. Animals from exposed ields seem to be in worst body conditions and in a deicitary energy balance when compared to animals from the reference locations. The results of the mesocosm study validated the correlative data obtained in the ield survey. While the laboratory approach showed that environmentally relevant dosages of chlorpyrifos can afect P. bocagei. According to our data, P. bocagei seems to be a suitable indicator of sub-lethal exposure to pesticides.

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The last catalogue of the Reptile collection of the Museu Bocage dates back to 1972. The collections were lost in a fire 25 years ago, so it was necessary to analyse the effort of various donors, who offered the Museum a new collection. The evaluation of the sampling effort is important to gather information (on quantity and origin) and the various samples of the species of lacertid lizards present in the Museum and enables an operational and practical visual imaged of that information. The obtained results prove that the study of the sampling effort using ordinary kriging method is robust and accurate on identifying gaps (under or no sampled areas) and a helpful tool to fulfil them. Although beginning to the representative of the Portuguese fauna, the lacertid collection of the Museu Bocage needs major improvements given that it still does not illustrate the actual distribution of lacertid lizards in Portugal. In order to stand for modern standards, when complementing the existing collection we must take into account not only the information provided by the analysis but also conservationist concerns (not killing animals) and the establishment of co-partnerships with congener institutions as well. Nowadays Museums have the duty, more than ever, to keep their collections well housed, cared for and accessible to the scientific community.

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Few studies have analyzed spatially the home ranges of small lizards, namely in the Iberian Peninsula. We studied the home ranges of two Iberian lacertids (Podarcis bocagei and P. hispanica) in a sympatric situation. Fieldwork was performed in a small area (Moledo beach, 0.8 ha) in the North-West of Portugal, situated within an urban area, limited by buildings, stone walls and small ields of traditional agriculture. The study area is characterized by four stones walls, and a small beach area with rocks and vegetation. Both species are in strict sympatry and present high densities. We captured 76 lizards of both species, and the exact position was geo-referenced with a professional Trimble GPS (horizontal error lower than 50 cm). We collected morphological measures, as well as tissue and blood samples; we marked the lizards with colored inks and release them in the same place of capture. We followed marked and unmarked lizards during seven days, making random paths around the stone walls and the beach. By visual contact and without recapture them, we recorded lizards’ positions with the GPS and collected other information such as species, sex, age, social interactions, environmental temperature, humidity, and substrate temperature. We calculated the home ranges of marked individuals with minimum convex polygons (MCP), within a Geographical Information System. We analyzedthe relationships between males and females of diferent species and their movements. We recorded a total of 774 individuals of Podarcis bocagei (339 females and 435 males), of which 236 were marked, and 243 individuals of Podarcis hispanica (126 females and 117 males), of which 72 were marked. Preliminary results showed movements among distant walls and overlapping of female home ranges with the home ranges of several males.

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The basic anatomy of the lacertid hemipenis (intromittent organ) and methods for its investigation are described. In many members of the Lacertidae, the hemipenis has a structure quite unlike that of other squamate reptiles: the distal lobes of the retracted organ are complexly folded and there is a well-defined supporting structure of dense connective tissue, the armature. This incorporates blood sinuses and has an intramuscular portion embedded in the m. retractor penis magnus and two club-shaped bodies, the clavulae, that support the lobes in the erect organ. Unarmatured hemipenes occur in some lacertids and, like those of other squamates, possess sac-like lobes in the retracted state, but they are singular in having the lobes invested by the m. retractor penis magnus. It is argued that many of these apparently primitive hemipenes are in fact secondary derivatives of the armatured type. There is considerable inter-specific variation in hemipenial structure which is described systematically. In some cases this involves differences in size, asymmetry and simplification, which may arise as physical isolating mechanisms and is useful in distinguishing otherwise very similar species, particularly in the genus Mesalina (p. 1253). Other shared derived hemipenial features provide useful information about relationships between species and higher taxa and a summary of the hypotheses that they support is given (p. 1254).

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Twenty-four species of lacertid lizards were examined at 31 sites in western Yogoslavia, Greece and Iberia. Comparative observations were made on over 4500 individual lizards, noting such features as times of activitiy, hunting methods, diet, micro- and macrohabitat, refuges used and body temperature. These data are used as a basis for assessing resource partition in related sympatric species, for find out whether different systematic groups have characteristic types of niche, and for comparing cumminity structures in the peninsulas of southern Europa. Acanthodactylus erythrurus, Algyroides marchi, Algyroides moreoticus, Algyroides nigropunctatus, Lacerta agilis, Lacerta graeca, Lacerta horvathi, Lacerta lepida, Lacerta monticola, Lacerta oxycephala, Lacerta mosorensis, Lacerta schreiberi, Lacerta trilineata, Lacerta viridis, Lacerta vivipara, Podarcis bocagei, Podarcis erhardii, Podarcis hispanica, Podarcis melisellensis, Podarcis milensis, Podarcis muralis, Podarcis peloponnesiaca, Podarcis sicula, Podarcis taurica, Psammodromus algirus, Psammodromus hispanicus.

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We studied morphological and molecular characters in Podarcis bocagei and P. hispanica lizards in the Ria de Arosa archipelago in coastal Galicia, Spain. Contrasting published information about insular distributions indicated that morphological species identification is problematic. Instead, we identified 145 lizards from 13 islands and several mainland populations by a panel of partially diagnostic nuclear protein loci and through the DNA sequencing of a stretch of 297 base pairs (bp) of the mitochondrial cytochrome-b gene. Correspondence between the molecular identifications was complete, with the exception of two lizards that carried P. hispanica mitochondrial DNA in a P. bocagei nuclear background. The combined results indicate past hybridization events and the oversea dispersal of a female lizard over a distance of about 500 m. Fourteen morphometric and 12 meristic characters were measured in 187 lizards from 15 islands and several mainland reference populations. Multivariate analysis revealed significant differences between sexes and between species. Two island populations for which no molecular data were available were identified as belonging to P. bocagei from morphology. Locally, P. bocagei and P. hispanica have an essentially parapatric distribution, with P. bocagei in the northeast and the islands of the inner Ria, and P. hispanica in the southwest and the islands of the outer Ria. Common patterns for island populations are an increase in absolute size, a decrease in three relative head width parameters, and a lower nuclear genetic heterozygosity than on the mainland. No correspondence was observed between morphological and molecular patterns of intra-specific differentiation. Also, linear distance between populations did not help to explain the results. Because the Ria de Arosa is a flooded river valley, the local bathymetry reflects the order in which the islands have become isolated from the mainland since the sea-level rise started c. 14,000 years ago. However, the temporal order of isolation was not associated with the pattern of population differentiation. The island populations of P. hispanica seem to have diverged over multiple genetic and morphological axes, with no obvious relationship to inter-population distance, coastline history, island ecology, or species habitat preferences. The results indicate that genetic drift is the main force driving in population divergence. Oversea dispersal, that is gene-flow, appears insufficient to counter the morphological differentiation between some island populations. Hypothesis on the biogeographical history of P. bocagei and P. hispanica in western Galicia are discussed.

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Archaeolacerta s. l., in the currently utilized sense results to be a polyphyletic taxon, composed of three monophyletic and phylogenetically well differentiated taxa: a)-Archaeolacerta (s. str.) Mertens, 1921, which includes three species: A. bedriagae, A. oxycephala, and A. mosorensis, distributed by the northern part of the Central Mediterranean (Corsica, Sardinia, and west Balcanic Peninsula). His most related genera seem to be Teira (his adelphotaxon in our study), Apathya, and Omanosaura. b)-Iberolacerta gen. nov., with two subgenera: Iberolacerta s. str. and Pyrenesaura subgen. nov. includes six species: I. monticola, I. cyreni, I. bonnali, I. aranica, I. aurelioi, and I. horvathi. They are distributed by the mountains of Western Europe (Iberian Peninsula, Pyrenees, Central and Eastern Alps and the north of the Dinaric Chains). His adelphotaxon are the Caucasian and Near East species of Darevskia gen. nov. c)-The third taxa is Darevskia gen. nov., which includes the species of the `L.` saxicola complex besides `L.` derjugini, `L.` praticola, and `L.` chlorogaster. They are distributed by Eastern Europe, Caucasus, and Near East. Other clear relationships among some taxa of the Eurasian Radiation [= `Eurasische linie` from (Mayer and Benyr, 1994)] have been found: Algyroides seems to be the sister group of the species considered of uncertain phylogenetic relationships belonging to the `L.` danfordi-laevis group and of Podarcis. `L.` brandtii seems to be a very primitive species within the Eurasian Radiation. Omanosaura and Apathya appear as sister taxa. `L.` graeca occupies a very basal position in the sister group of Podarcis and relatives (see above). `L.` andreanskyi is the sister species of the Darevskia nov. and Iberolacerta nov. clade. The assimilation of `L.` andreanskyi to Teira is very problematic. Also the `L.` parva group seems to be related to Timon. Some groups of species like `L.` brandtii, `L.` parva-fraasii and `L.` danfordi-laevis probably merit generic rank, but it is necessary a more deep study before to take a decision.

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Arboreal behaviour in Podarcis bocagei inhabiting a coastal habitat in Northwestern Spain is reported in this note. Ornamental olive trees were selected as a refuge rather than as bask-ing areas.

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On a herpetological trip to the north-west of the Iberian Peninsula, the authors were able to record a total of ten species of lacertid lizards along with other reptile species. In particular, the densely populated Portuguese Atlantic coast proved to be a species-rich lizard habitat.

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We measured the level of fluctuating asymmetry (FA) in head shape, head scalation and femoral pores in two lizard species (Podarcis bocagei and Podarcis hispanica) from 13 islands and 15 mainland localities in the Ria de Arosa archipelago of north-western Spain. Given the recent geological history of the region, the degree of isolation to which lizard populations have been subjected can be ordered along a spatio-temporal gradient, yielding the following hypotheses to be tested: FA will be higher (1) in island populations than in mainland populations; (2) on remote islands than on islands close to the mainland; (3) on small islands than on large islands. Molecular genetic data suggest that P. hispanica is autochthonous in the Ria de Arosa, whereas P. bocagei is a more recent arrival. Therefore, we predict also (4) a higher level of FA in P. hispanica than in P. bocagei. Statistically significant results were obtained for head-shape asymmetry, supporting the second and the fourth hypotheses. With an overall meristic asymmetry index, none of the hypotheses were corroborated, whereas for certain independent meristic traits, the first, the third and the fourth hypotheses were partially supported. Both head shape and meristic traits constitute precise measures of FA, but FA is more convincingly expressed in head shape and in single meristic traits than in overall meristic traits asymmetry. We conclude that FA reflects population isolation and may be a good indicator of developmental instability. It seems worthwhile to test for FA in a landlocked system under environmental and genetic stress, for the purpose of conservation biological assessments.

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During the month of June 1980 the authors observed several species of amphibians and reptiles in Portugal. Notes are given on their habitat. Problems concerning the determination of some species of the genus Podarcis are discussed.

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We carried out a morphometric study of the skull throughout postnatal ontogeny in 14 species of Iberian lacertid lizards belonging to the genera Acanthodactylus, Algyroides, Psammodromus, Podarcis, Zootoca, Timon and Lacerta s. latu with the aim of elaborating a key of identification for the studies species. A total of sixty one characters showing interspecific variation are identified. The characters are defined in articulated and disarticulated skulls.

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Intra- und interspecific variation are analysed fpr the post-natal skull of the lacertid lizards Gallotia galloti, Actnhodactylus erythrurus, Algyroides marchi, Lacerta monticola, Lacerta vivipara, Podarcis bocagei, Podarcis hispanica, Podarcis muralis, Psammodromus algirus and Psammodromus hispanicus. Individual variations identified include: the prsence/absence, number and morphology of some structures, mainly in the lacrimal, sclerotic ossicles and macilla; the number and location of foramina and the degree of ossification of some processes in chondrocranial bones. No differences bones. No differences have been found between the sexes with respect to presence and morphology of the bones but some species show sexual dimorphism in the size and robustness of the head and this may be reflected in the individual. The most substantial variations seen are ontogenetic: the appearances of new traits; development of articulations; differences between species in the timing of the stages of development of a given structure; changes in the relative position of some cranial elements; and the degree of calcification or ossification of processes. A total of 63 characters showing interspecific variations are identified. Some have not previously been described while others used in previous studies are redefined here on the basis of new morphological information obtained.

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Males of many lizard species have longer tails than similarly-sized females. We hypothesized that this dimorphism is induced by a longer non-autotomous tail part in males, which is associated with the presence of the copulatory organs at the tail base, and presumably reduces the males` ability to escape predation by tail shedding. A compensatory mechanism would be an increase of total tail length in males, to achieve equal lengths of the autotomous tail part in both sexes. A critical prediction of this `morphological constraint` hypothesis is that the extent of dimorphism in total tail length increases with the magnitude of sexual differences in length of the non-autotomous tail base. We tested this prediction through a comparative study in a small clade of lacertid lizards. Within each of nine species, sexual differences in length of the non-autotomous tail base and in total tail length do not change with body size. All species, except one, exhibit a clear male-biased dimorphism in length of the non-breakable tail base. In all species studied, males have longer tails than females. We used the method of phylogenetically independent contrasts to explore the interspecific relation between dimorphism in length of the tail base and sexual differences in total tail length. Contrary to our prediction, we found no evidence for a positive correlation between the extent of dimorphism in both traits. Thus, constraints imposed by the male copulatory organs on tail autotomy do not seem to be a significant factor in the evolution of dimorphism in tail length in this clade of lacertid lizards.

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In many animals, chemical signals play an important role in species recognition and may contribute to reproductive isolation and speciation. The Iberian lizards of the genus Podarcis, with up to nine currently recognized lineages that are often sympatric, are highly chemosensory and provide an excellent model for the study of chemically mediated species recognition in closely related taxa. In this study, we tested the ability of male and female lizards of two sister species with widely overlapping distribution ranges (Podarcis bocagei and P. hispanica type 1) to discriminate between conspecific and heterospecific mates by using only substrate-borne chemical cues. We scored the number of tongue flicks directed at the paper substrate by each individual in a terrarium previously occupied by a conspecific or a heterospecific lizard of the opposite sex. Results show that males of P. bocagei and P. hispanica type 1 are capable of discriminating chemically between conspecifics and heterospecifics of the opposite sex, but females are not. These results suggest that differences in female, but not male, chemical cues may underlie species recognition and contribute to reproductive isolation in these species. The apparent inability of females to discriminate conspecific from heterospecific males, which is not because of reduced baseline exploration rates, is discussed in the context of sexual selection theory and species discrimination.

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To gain insight into the role of behaviour in promoting reproductive isolation between Podarcis bocagei and Podarcis morphotype 1, sister species belonging to the P. hispanica species complex, we conducted field and laboratory observations. We studied lizards in rocky outcrops and man-made stone walls in Moledo do Minho (NW Portugal), where these two species occur in syntopy. During the reproductive season of 2006 we video-recorded interactions between free-ranging individuals in a stone wall ca 50 m long. These observations allowed us to document and describe, in natural conditions, the behaviours exhibited during male-female interactions. Interactions were assigned one of the following scores, reflecting increasing levels of courtship behaviour: (1) No response: either lizards remain close or one of them moved away; (2) Displays: one of the lizards exhibited visual displays (e.g. foot-shakes strut) or the male followed the female; (3) The male bites the female tail; (4) The male attempts (abdominal bite) or copulates with female. A comparison of intra- and interspecific interactions revealed that the former received significantly higher scores, indicating that lizards exhibited more courtship towards conspecifics. During the spring of 2008 we used an experimental approach to assess, now under controlled laboratory conditions, whether intraspecific encounters would result in more courtship (i.e. higher courtship scores) than interspecific encounters. Briefly, we staged male- female encounters in glass terraria and video-recorded ensuing interactions for 30 min. Analyses revealed that interaction scores were again significantly higher in intraspecific encounters, and also that males displayed significantly more in the presence of conspecific females. These results suggest that males are able to discriminate conspecific from heterospecific females and stress the importance of behavioural mechanisms in maintaining reproductive isolation in this complex group of lacertids. The study of reproductive interactions within this group adds an important new dimension to ongoing research about their phylogeography.

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The present work studies the herpetological community of Caural (NW, of Spain): biogeography and ecology. Biogeography altitudinal distribution of species, altitudinal variation in abundance of species and stratification reflect a decrease with the altitude bound to climatic reasons and reproductivity in reptiles and to lesser heterogeneity of biotopes in amphibians. There is a greater altitudinal stratification of the subcommunity of amphibians than of reptiles. Ecology: the different physiological requirements of amphibians from land habitats and lizards are reflected in a temporal alternation (seasonal and daily) in the exploitation of the same trophical resource. The snakes occupy anoth trophical level (predators and superpredators) and the tritons live inanother habitat (subaquatic) with different resources and they have a different diet. The problem of interspecific competition is dealt with in two concrete cases: Lacertidae and Triturus. Great differences in size or corporal shape and behaviour permit coexistence but in the opposite case they separate themselves throughout the area, distributing the biotopes according to their physiological preferences. There are cases (Lacerta schreiberi and Triturus boscai) in which there appears to be a spatial displacement due to a diffuse competition with congenital species. One highlights the importance of the habitat in the food diet of some species, which becomes very clear comparing the tritons with the lizards and amphibians from land habitats. One questions the problem of stability and heterogeneity (climatic and spatial), competition and predation as factors which impel and influence the evolution of communities and species. In relation with studies realized in other areas we see that the herpetological community of Caurel represents the transition from the Mediterranean region to the European shore region, very much in accordance with the climatic and botanical characteristics of the zone, where the seasonality plays an important role just as the glaciarism appears to have determined a reptile community evolutionally young.

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In this work the actual state of knowledge on biogeography of amphibians and reptiles from Galicia is exposed in a synthetical way based upon distributional data gathered during the last 10 years. The geographic distribution of species and communities on latitudinal and altitudinal gradients indicates that climate and biotope are more important factors than latitude or altitude over sea. The presence of relict populations of eurosibirian taxa and the geographic distribution of endemism poit to the importance of climatic alterations during the Quaternary Age (Glaciarism). Thus, the eurosibirian taxa were driven towards Galicia following both the Cornice and the Cantabrian Chain as a mountains, giving rise to sub-especiation or speciation in allopatry of some endemisms. As the present data are poor we must consider our conclusions as a first hypothesis.

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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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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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Pesticide exposure has been related with thyroid disrupting effects in different vertebrate species. However, very little is known about the effects of these compounds in reptiles. In the Mediterranean area, lacertid lizards are the most abundant vertebrate group in agroecosystems, and have been identified as potential model species for reptile ecotoxicology. The aim of this study was to understand if the herbicides applied in corn fields have thyroid disruptive effects in the lizard Podarcis bocagei. Adult male lizards were captured in north-western Portugal in corn fields treated with herbicides (exposed sites), and in organic agricultural fields (reference sites). Thyroid and male gonad morphology and functionality, and testosterone levels were investigated through histological, immunohistochemical and biochemical techniques. Lizards from exposed locations displayed thyroid follicular lumens with more reabsorption vacuoles and significantly larger follicular area than those from reference fields. Furthermore, testes of lizards from exposed locations had significantly larger seminiferous tubule diameters, significantly higher number of spermatogenic layers and displayed an up-regulation of thyroid hormone receptors when compared with lizards from reference areas. These findings strongly suggest that the complex mixture of herbicides that lizards are exposed to in agricultural areas have thyroid disrupting effects which ultimately affect the male reproductive system. Alachlor, which has demonstrated thyroid effects in mammals, may be largely responsible for the observed effects.

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The interspecific relationships among female size, clutch size, egg size, and hatchling size were examined for 64 European lacertids. The eggs of all species increased linearly in both linear dimensions and mass during incubation. Across species initial egg mass was positively correlated with juvenile mass, with an allometric relationship exponent of 0.87. Initial egg mass across species increased proportionally with female mass to the power of 0.57. Moreover, an increase in maternal mass was also accompanied by an increase in clutch size. The number of eggs per clutch across species scaled with female mass to the power of 0.39. Removing the effect of female mass resulted in a negative correlation between egg mass and clutch size. Species for which the average egg size was lower than expected on the basis of female mass, tended to have larger relative clutch size. The total egg mass per clutch was about one third of female mass (exponent 0.94).

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Die Arbeit basiert auf der Messung verschiedener metrischer Charaktere (Kopf-Rumpf-Länge, Kopflänge, maximale Kopfhöhe, maximale Kopfbreite und Rumpf-Länge) an insgesamt knapp 500 nordspanischen Eidechsen. In die Analyse gingen Mesungen an Podarcis bocagei, Podarcis hispanica, Podarcis muralis, Lacerta monticola, Lacerta vivipara, Lacerta lepida, Lacerta schreiberi, Lacerta viridis (=L. bilineata) ein. Demnach ist das ontogenetische Wachstum von Kopf und Rumpf bei männlichen Tieren isometrisch, während Weibchen hingegen ein allometrisches Wachstum des Rumpfes aufweisen. Mit der verwendeten Methodik (Felsenstein) ließ sich der Geschlechtsdimorphismus am besten durch den weiblichen Fortpflanzungsaufwandes (u. a. Gelegegröße) und damit durch die überproportionale Rumpflänge weiblicher Tiere erklären. Bei Arten mit geringer Fruchtbarkeit bzw. konstanter Gelegegröße (hier: Podarcis hispanica) besteht hingegen nur ein geringer Selektionsdruck in Richtung auf große Weibchen. Dann können Männchen zum größten Geschlecht werden. Ebenfalls durchgeführte Größenmessungen an Beutetieren (Magen-Darminhalte) ergaben vergleichsweise gering abweichende Einnischungen männlicher und weiblicher Eidechsen. Eine Analyse der Mechanismen, die zur Ausprägung von Geschlechtsdimorphismen beitragen, ist sicherlich verdienstvoll. Allerdings ist fraglich, ob die hier verwendeten Methoden allein zu einer befriedigenden Klärung führen. Zum einen wurden pro Art nur wenige Individuen (zwischen 22 und 90) untersucht, was - auch nach Ansicht des Autors - statistisch unbefriedigend ist. Wichtiger dürfte aber wohl sein, dass keinerlei Hinweise darauf vorliegen, inwieweit die aus einem relativ begrenzten Gebiet stammenden Freilandfänge als “typisch“ für die Art gelten können. Hinzu kommt, dass sich mit der verwendeten Methode nur die Parameter weiblicher Fortpflanzungsaufwand und unterschiedliche Nahrungseinnischung, nicht jedoch der dritte vom Autor angesprochene Parameter “Kämpfe zwischen den Männchen“ untersuchen lässt. Zumindest in diesem Kontext bedarf es weiterer Daten, die nur durch Verhaltensbeobachtungen lebender Eidechsen zu gewinnen sind.

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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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Unravelling when divergent lineages constitute distinct species can be challenging, particularly in complex scenarios combining cryptic diversity and phylogenetic discordances between different types of molecular markers. Combining a phylogenetic approach with the study of contact zones can help to overcome such difficulties. The Podarcis hispanicus species complex has proven to be prosperous in independent evolutionary units, sometimes associated with cryptic diversity. Previous studies have revealed that one of the species of this complex, P. guadarramae, comprises two deeply divergent yet morphologically indistinguishable evolutionary units, currently regarded as subspecies (P. g. guadarramae and P. g. lusitanicus). In this study we used molecular data to address the systematics of the two lineages of Podarcis guadarramae and the closely related P. bocagei. Firstly, we reconstructed the species tree of these three and two additional taxa based on 30 nuclear loci using the multispecies coalescent with and without gene flow. Secondly, we used SNPs obtained from RADseq data to analyze the population structure across the distribution limits P. g. lusitanicus and P. g. guadarramae, and for comparison, a contact zone between P. bocagei and P. g. lusitanicus. Nuclear phylogenetic relationships between these three taxa are clearly difficult to determine due to the influence of gene flow, but our results give little support to the monophyly of P. guadarramae, potentially due to a nearly simultaneous divergence between them. Genetic structure and geographic cline analysis revealed that the two lineages of P. guadarramae replace each other abruptly across the sampled region and that gene flow is geographically restricted, implying the existence of strong reproductive isolation. Podarcis bocagei and P. g. lusitanicus show a similar degree of genetic differentiation and reproductive isolation, with very low levels of admixture in syntopy. These results support that all three forms are equally differentiated and reproductively isolated. In consequence, we conclude that the two former subspecies of Podarcis guadarramae constitute valid, yet cryptic species, that should be referred to as P. lusitanicus and P. guadarramae.

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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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Climate change has been widely accepted as one of the major threats for global biodiversity and understanding its potential effects on species distribution is crucial to optimise conservation planning in future scenarios under global change. Integrating detailed climatic data across spatial and temporal scales into species distribution modelling can help to predict potential changes in biodiversity. Consequently, this type of data can be useful for developing efficient biodiversity management and conservation planning. The provision of such data becomes even more important in highly biodiverse regions, currently suffering from climatic and landscape changes. The Transboundary Biosphere Reserve of Meseta Ibérica (BRMI; Portugal-Spain) is one of the most relevant reserves for wildlife in Europe. This highly diverse region is of great ecological and socio-economical interest, suffering from synergistic processes of rural land abandonment and climatic instabilities that currently threaten local biodiversity. Aiming to optimise conservation planning in the Reserve, we provide a complete dataset of historical and future climate models (1 x 1 km) for the BRMI, used to build a series of distribution models for 207 vertebrate species. These models are projected for 2050 under two climate change scenarios. The climatic suitability of 52% and 57% of the species are predicted to decrease under the intermediate and extreme climatic scenarios, respectively. These models constitute framework data for improving local conservation planning in the Reserve, which should be further supported by implementing climate and land-use change factors to increase the accuracy of future predictions of species distributions in the study area.

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Lacertids are the dominant group of lizards throughout the Mediterranean Basin. Their role in food web transfer of matter and energy from arthropods and other small invertebrates to birds and mammals constitutes a major function within Mediterranean ecosystems. For many years, prey consumption by lacertids was thought to be almost indiscriminate, not much more than a byproduct of habitat use. However, increasing evidence does not support this passive view. Analyses of prey availability have revealed active prey selection/avoidance in several species. Others show an internal tendency (i.e., historical constraints) to consume specific animal items (ants, clumped prey) or plant matter (seeds, nectar, pollen, leaves). Behavioural experiments showed that lacertids not only identify different prey types by both visual and chemical cues but also modify their feeding behaviour integrating past experiences. Furthermore, size, sex, reproductive state, body condition, tail loss and probably other lizard features are relevant for feeding ecology. However, less attention has been devoted to abiotic factors such as temperature and humidity. More experimental studies of the influences of competitors, predators and parasites on diet are needed. Even though it is controversial, optimal foraging theory provides a conceptual background for future studies. The evolutionary history of the various lacertid lineages, which constrains their morphology and physiology and eventually produces exaptative traits, is to be considered as well. Finally, methodology in field sampling, lab work and statistical analysis needs to be developed. Recommendations are given as to when and where to sample, which compartment should be analysed, which is the appropriate sample size, how to assess trophic availability, which statistical descriptors should be used and how they should be compared.

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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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Lacertids are considered conservative in terms of their thermal physiology. Namely, whereas their temperatures in the field vary according to the environment, their preferred body temperatures (Tp) when free of thermal constraints remain similar within species even if under different climate regimes. In contrast, phylogenetically distant lacertid lineages sharing the same habitats may display distinct Tps more related with their biogeographic origins. The evolutionary limits of this so-called thermal rigidity remain, nevertheless, undefined. Here, the variation of Tp is analysed across the Iberomaghrebian Podarcis clade (P. bocagei, P. carbonelli, P. vaucheri and P. hispanica sensu lato). For this monophyletic group of closely related forms, some still not formally described, the knowledge of their phylogenetic relationships, divergence times and palaeoecological scenarios is now considerably robust. A total of 229 adult lizards belonging to 11 lineages were exposed to a thermal gradient and temperatures were recorded at 9 time intervals within the period of activity. Once the effects of diel rhythms, size and pregnancy were excluded, variation in Tp was recorded between some of the forms. Lineages diverging during the Pleistocene displayed similar Tps whereas sister taxa separated since the Pliocene show significant differences even if current ranges overlap. The most basal lineages in the phylogeny selected for different temperatures even if sister taxa but approached others not directly related to them. Apparently, divergence times higher than 3 mya are able to promote Tp shifts in this species complex. Such results are discussed in relation with the evolutionary history of this group.

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The spermatogenic cycles of Podarcis bocagei and P. carbonelli, two lacertids previously considered conspecific, were analysed in northwest Portugal. Monthly sampling was carried out at three sites: a zone of strict syntopy and two sites of allopatry for both species, all of them geographically close and harbouring similar habitats. The maritime Atlantic climate, which allows males to remain active most of the year, plays a determinant role on the variation of the mixed- type cycle, typical of lacertids in temperate regions. Comparison with other populations and species indicates that climate modifies the pattern of spermatogenesis by restricting the beginning (but not the end) of the activity period. Results also corroborate previous studies that report an agreement between the variations in the gonads observed at both microscopic and macroscopic scales in the reproductive period. Peaks for spermatozoa were delayed in sympatry but competition cannot be directly concluded. Furthermore, relationships between the number of spermatozoa and body size suggest sperm competition and possibly a reproductive interaction between the species.

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A parasitological survey was carried out to determine the relationships between the helminth fauna and biological traits of the hosts in three Algerian populations of Podarcis lizards belonging to two different evolutionary lineages. Size, sex, and locality of collection, as well as the infracommunities and component communities, were analyzed. Very low values of parasite infection parameters and diversity were found in all three populations. This is in accordance with the feeding habits of these lizard hosts, which only eat animal prey and no plant matter. Spauligodon saxicolae (Nematoda: Pharyngodonidae), a specialist in lizards, is reported here for the first time in Africa. This nematode was the dominant parasite species for the three populations of lizards regardless their phylogenetic lineage, size, sex, and environmental conditions.

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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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Dans ce (ravail nous faisons une synthese de quelques données que nous avons obtenu sur la distribution et I`écologie de I`herpétofaune portugaise. Naus faisons aussi référence à la fréquence de capture de ces animaux pendan! les différents mois de I`année et à leur occurrence dans les principales montagnes du pays.

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Species that are able to solve novel problems through social learning from either a conspecific or a heterospecific may gain a significant advantage in new environments. We tested the ability of a highly successful invasive species, the Italian wall lizard Podarcis sicula, to solve a novel foraging task when social information was available from both a conspecific and an unfamiliar heterospecific (Podarcis bocagei). We found that Italian wall lizards that had access to social information made fewer errors, regardless of whether the demonstrator was a conspecific or a heterospecific, compared to Italian wall lizards that individually learnt the same task. We suggest that social learning could be a previously underappreciated, advantageous mechanism facilitating invasions.

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Interactions between lizards may differ depending on whether they encounter conspecifics or different species, and agonistic behavior and tolerance have been the subject of many studies (BRATTSTROM 1982, CARPENTER 1960, CREWS & FITZGERALD 1980, GRASSMAN& CREWS 1987, MILSTEAD 1957, STAMPS & TANA- KA 1981). Podarcis bocagei (Lacertilia: Lacertidae) is a common arboreal/saxico- lous lizard in north central Portugal. Males and females are sexually dimorphic. Cnemidophorus velox (Teiidae) is a terrestrial, all-female, triploid, parthenogenetic lizard found in deserts at relatively high altitudes in New Mexico, USA Both species are widely foraging insectivores, however, P. bocagei is territorial and has extended foraging times both daily and seasonally (GiL et al. 1988) whereas C. velox is not territorial and is active briefly during the day in the summer (BowKER et al. 1986). Both species frequently encounter conspecifics in nature. Here we examine social interactions occurring when conspecific individuals meet under laboratory conditions designed to encourage social interactions. The overall goals are to determine if a dominance hierarchy exists, to determine the intensity of intraspecific interactions, and finally to establish correlates with dominance and aggressive/submissive behavior.

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Food availability and diet composition were examined in sympatric populations of Lacerta schreiberi and Podarcis bocagei at a locality in the Cantabrian mountains (province of León, Spain). Neither species uses trophic resources in proportion to their relative abundance. Extensive overlap in diet composition has been found between P. bocagei and juvenile L. schreiberi. Although both species feed mainly on spiders, juvenile L. schreiberi eat the most abundant and P. bocagei select larger prey sizes. Subadult and adult L. schreiberi select beetles and grasshoppers.

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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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Maintaining body temperature is essential for the optimal performance of physiological functions. Ectotherms depend on external heat sources to thermoregulate. However, thermoregulation may be constrained by body condition and hydration state. Autotomy (i.e., the voluntary shed of a body part) evolved in various animal lineages and allowed surviving certain events (such as predator attacks), but it may affect body condition and volume/surface ratios, increase dehydration and constrain thermoregulation. In the framework of a general analysis of the evolution of autotomy, here we assessed the effects of tail loss on the thermal preferences and evaporative water loss rates (EWL) in the lizard Podarcis bocagei, integrating the thermal and hydric factors. We did not observe shifts in the thermal preferences of experimentally autotomized lizards when compared to the controls, which contradicted the hypothesis that they would raise preferred temperature to increase metabolic rates and accelerate regeneration. Evaporative water loss rates were also similar for tailed and tailless in dividuals, suggesting negligible increase of water loss through the injury and no specific ecophysiological re sponses after autotomy. Therefore, the changes observed in autotomized lizards in the field are to be considered primarily behavioural, rather than physiological, and thermoregulation could be secondarily affected by behavioural compensations for an increased predation risk after autotomy. Functional studies are necessary to understand how lizards’ interaction with the environment is altered after autotomy, and further studies including different dehydration levels would be useful to fully understand the effect of water shortage on lizards’ per formance after caudal autotomy.

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Fire has impact on reptile communities with marked shifts in community composition between burnt and unburnt areas. These shifts are often related to the preference of reptile species throughout early or late post-fire successional habitats. Areas located in transition zones between bioregions harbor complex reptile communities with a mixture of biogeographic affinities. In these biogeographic crossroads, since fire simplifies the habitat structure, we expected simpler (lower alpha diversity) and more similar (lower beta diversity) reptile communities within burnt than within unburnt localities. We have tested this hypothesis in a transition zone between the Atlantic and Mediterranean bioregions in northern Portugal. Reptiles were surveyed in five localities (8 times per locality) along fire edges in which each locality was composed of one burnt and one unburnt transect. In total, 588 reptiles from 10 species were recorded. Unburnt transects had higher alpha and beta diversity and higher relative abundance of non-Mediterranean individuals than did burnt transects. A redundancy analysis also showed contrasting responses of Mediterranean and non-Mediterranean species, the former increasing and the latter decreasing after fire. Our study demonstrates that fire reduced the complexity of the reptile community, with benefits towards Mediterranean species due to its environmental preferences and long evolutionary association to fire. In biogeographic crossroads such as the study area, the retention of long unburnt vegetation is expected to maintain more diverse reptile communities.

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Transition zones between bioregions can sustain the coexistence of species with different biogeographic affinities through ecological niche segregation at a finer scale. Reptiles can serve as a good model group to evaluate the mechanisms behind such coexistence patterns especially due to their thermal sensibility to microhabitat structure. We examined whether reptile species spatially segregate in their microhabitat use and whether this segregation reflects the biogeographic affinities of species. Reptiles were surveyed across a 1.5-km transect located in the Peneda-Gerês National Park. We used 478 GPS observations of nine sympatric non-Mediterranean and Mediterranean species collected during field surveys in 2012 and 2013. Each observation was assigned to a microhabitat-openness measurement. Differences in microhabitat selection between Mediterranean and non-Mediterranean species were studied with factorial ANOVA analyses. The similarity in microhabitat selection between pairs of species was compared with the similarity in their Iberian distributions by a Partial Mantel test checking for the genetic distances between species pairs in a phylogenetic framework. Comparing a random selection of points along transect, we found that non-Mediterranean species used more closed microhabitats than Mediterranean species. No differences in this pattern were found between snakes and lizards. The Mantel test showed that microhabitat selection was correlated with the Iberian geographic distributional patterns of species. The complex vegetation structure of this area is most likely responsible for the availability of different microhabitats, which promotes high reptile species richness and governs their sympatric coexistence at a finer scale.

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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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Se ha estudiado la incidencia de infecciones externas en una población noroccidental ibérica del lacértido Podarcis bocagei. Estas infecciones llegaron a afectar al 20-30% de los individuos observados en el invierno de 1989-1990 y al 40-60% en el invierno 1990-1991. Los porcentajes de individuos afectados fueron mayores en los meses más lluviosos y fríos. No se encontraron diferencias en las tasas de recaptura entre los individuos afectados y no afectados, lo que parece indicar que estas afecciones no tienen un efecto directo detectable en las tasas de mortalidad. La presencia de esta enfermedad afectó a la reproducción: el 28% de las hembras adultas de talla grande (LCC > 55 mm) que habían estado infectadas, no participaron en el primer período de puestas, durante mayo, mientras que tan sólo el 2.2% de las no afectadas de la misma talla no se reprodujo en ese período. Los individuos jóvenes afectados mostraron tasas de crecimiento significativamente menores que los ejemplares de su misma edad no afectados. Por lo tanto, el efecto de estas infecciones sobre el éxito reproductor se manifiesta principalmente afectando a la participación en la reproducción de las hembras adultas y a las tasas de crecimiento de los individuos jóvenes.

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This paper presents data on reproductive and fat body cycles of the oviparous lacertid lizard Podarcis bocagei in northwest Spain. Testes exhibited their maximal volume during December – March and decreased in size throughout the summer. This pattern agrees with the “mixed type” of spermatogenetic cycle proposed by Saint Girtons (1963, 1982). Vitellogenesis started at the end of March or April. Females with oviductal eggs were found from mid-April to mid-July. Oviposition occurs between mid-May and the end of July. Fat bodies of males were smallest during the spring mating period. Female fat body volumes declined during vitellogenesis. No siognificant decline of lipid stores was detected during the winter period (October – March).

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This paper presents data on aspects of the reproductive ecology of a population of Podarcn hocagei in northwestern Spam, as monitored over a two-year penod (1990-1991) Data were obtained pnncipally on the basis of mark-recapture expenments, but also from laboratory hatching studies Mating took place between the end of March and July Dunng the laying penod, from May to July, 8 5% of reproductive females produced three clutches, 52 1% two clutches, and 39 4% one clutch In general, single clutches were produced by small females Only a small proportion of large females produced three clutches Mean clutch size was 4 8 eggs (range 4-7) in May, 4 3 (2-6) in June and 3 9 (2-4) in July There was sigmficant vanauon in the mean snout-to-vent length (SVL) of females laying in each month of the season Both clutch size and mean single-egg volume increased with mother`s SVL There was a significant partial correlation between egg volume and clutch size when both mother`s SVL and month of laying were held constant There was no significant between-year vanation in clutch size, breeding females` SVL, egg weight or relative clutch mass A delay in the timing of reproducuve events in one year (1991) is attnbutable to adverse weather conditions dunng early spnng Hatching occurred between July and September Hatch success (as estimated in 1989, 1990 and 1991 from natural nests at the study site) was high, ranging from 83% in 1991 to 91% in 1989 The mean SVL of female hatchlings was greater than that of male hatchlings By contrast, adult females had lower mean SVL than adult males

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Colonization by reptiles and amphibians of the spoil benches of the Meirama opencast lignite mine in northwest Spain was studied over the 10 years following the start of the revegetation process. At Meirama, spoil benches are initially fertilized and hydroseeded with a pasture mix, but are subject to little subsequent management and are gradually colonized by scrub vegetation characteristic of the region. Herpetofauna censuses were carried out (1) yearly on a single 2 ha plot over the 6 years following hydroseeding, and (2) in a single year on ten 2 ha plots which had been hydroseeded between 0 and 10 years previously. In addition, censuses were carried out on three undisturbed `control` plots close to the mine. Fifteen species (nine amphibians and six reptiles) were detected in one or more of the spoil-bench plots. Recolonization is particularly rapid after the first 2-3 years posthydroseeding, and is clearly closely related to habitat development due to revegetation and natural succession. The first reptile species to colonize the study plots was Podarcis bocagei (first year), and the first amphibians Alytes obstetricans and Rana perezi (second year). Species composition was most similar to that in control plots in the oldest (ten-year-old) spoil plots.

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En el presente artículo se realiza un inventario de las especies de anfibios y reptiles presentes en las islas de Galicia. También se presentan algunos datos sobre la ecología y la morfología de algunas poblaciones insulares de estas especies. La herpetofauna de las islas gallegas está compuesta por tres especies de Urodelos, siete de Anuros, siete de Saurios y cuatro de Ofidios. Las especies más ampliamente distribuidas en las islas de la costa de Galicia son las lagartijas del género Podareis. Por otro lado, las especies de anfibios son las que tienen una distribución más restringida, no habitando más que las islas de mayor tamaño, con la excepción de algunas poblaciones de Salamandra salamandra, que viven en pequeños islotes.

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Adults of Bocage`s Wall Lizard (Podarcis bocagei), a lacertid endemic to the northwestern Iberian Peninsula, generally show marked sexual dichromatism with males having green dorsal coloration and females brown dorsal coloration. In some populations, however, some adult females show dorsal coloration similar to that of males. I used mark-recapture methods to study a population of this type on a coastal promontory in A Coruña. A proportion of adult females showed green dorsal coloration during the reproductive period (April–July), but not during the rest of the year. Green dorsal coloration was most frequent among females with oviductal eggs, whereas preovulatory females with vitellogenic follicles in all cases showed brown dorsal coloration. These findings suggest that the green dorsal coloration observed in some females may be related to stage in the reproductive cycle, with green coloration developing during the gravid phase (i.e., with fertilized eggs). The findings also raise the possibility that the function of this coloration may be to avoid harassment and copulation attempts by males following fertilization. According to this hypothesis, females show brown dorsal coloration during the receptivity period (and during the nonreproductive period). Following fertilization, and possibly as a result of the hormonal changes that accompany ovulation and fertilization, they develop green dorsal coloration. This suggests that coloration, in these lacertids as in other reptiles, plays an important role in social communication.

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Mark-recapture techniques were used to investigate population size, age distribution, size distribution and sex ratio in a population of the lacertid lizard Podarcis bocagei in an abandoned gravel pit in northwest Spain. The study was carried out over a 2-year period. Despite relatively high maximum longevity, the population age distribution was characteristic of small, short-lived lizard species (i.e., there was a relatively high proportion of immature individuals).Population size declined over the study period, largely because of a drop in the number of immature animals: this may be partially attributable to density-dependent factors, but was probably due largely to a decline in habitat favourability as a result of colonisation of the study site by vegetation. The sex ratio was signi. Cantly female-biased in all cohorts studied, not only among adults but also among juveniles and sub-adults. However, sex ratio at hatching (as investigated by laboratory hatching of clutches laid by captured pregnant females) did not differ signi. cantly from one-to-one. There was no difference found in survival probabilities between males and females. The observed bias in sex ratio must therefore be attributed to between-sex differences in net emigration.

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The main threats to the amphibians and reptiles of Galicia (NW Spain) are studied in relation to the information obtained during the last 30 years (1975-2005). The most negative factors affecting amphibians and reptiles in this territory are the human alteration of both aquatic and terrestrial habitats, and the introduction of exotic species (mainly freshwater crustaceans and fishes). These menaces have led to the extinction of some Galician populations of amphibian and reptiles. Other factors, such as emergent diseases, can be very important, but they have not been studied yet. This last threat is one of the main causes for the global amphibian decline. The recent status of the Galician herpetofauna is also analysed. According to the main population declines and areal reduction observed, the most endangered amphibian species is Pelobates cultripes. Other amphibians, such as Chioglossa lusitanica, Rana iberica and Rana temporaria, are also importantly put at risk. The most endangered reptile species is Emys orbicularis. The restricted and limited Galician populations of Lacerta vivipara, Chalcides bedriagai and Vipera latasti are also threatened. The available information for other reptile species (Blanus cinereus, Psammodromus hispanicus, etc.), is still very scarce, but it is possible that their situation is also precarious. The particular status of the insular populations of amphibians and reptiles from the islands off the Galician coast is also analysed.

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Changes in the coloration of the lacertid lizards Iberolacerta monticola and Podarcis bocagei with age in populations from NW Spain are described. The onset of sexual maturity in P. bocagei males involves a change in the ventral (yellow) and dorsal (green) colorations, which is different from immature males (dorsally brownish in color). In I. monticola males, the ventral coloration also changes to a deep green when they reach maturity, while the dorsal coloration remains brownish as in the immature specimens. In this species, the green dorsal coloration is acquired gradually after maturity. Only the oldest individuals have a predominantly green dorsal coloration. The differences between the two species in the time males take to acquire the green dorsal coloration could be related to their diff erent longevity. The coloring is acquired gradually in the most long-lived species ( I. monticola ). A field study was carried out on the behaviour of adult males of I. monticola during the reproductive period. The males with green dorsal coloration were seen to pair with females signifi cantly more frequently than those with the brownish dorsal color. The increase in the green dorsal coloration (conspicuous) with the size and age of the males of this species would appear to have a clear function as an intersexual or intrasexual signal.

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Tail tip removal is a common method for obtaining tissue samples for genetics and other studies on lizards. This study evaluates the effect of tail tip autotomy on microhabitat selection in the lacertid Podarcis bocagei. Different-length tail fragments were experimentally removed from lizards of a small population. Forcing lizards to autotomise small tail tips (<1 cm) did not affect microhabitat selection. In contrast, a significant negative effect was observed in those lizards which underwent induced autotomy of the entire tail (> 5 cm). After autotomy these lizards were observed to favour more closed habitats, where predator avoidance is expected to be more efficient, although of potentially lower thermal quality.

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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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Múltiples trabajos sobre comunicación química han revelado que el sistema químiosensorial influye numerosos procesos subyacentes a la ecología y evolución de los reptiles. Sin embargo, sabemos muy poco sobre cómo la señalización química se ve influida por factores ecológicos y evolutivos. Sobre esta base, hemos elegido los lagartos como modelo de estudio para investigar la comunicación química, utilizando diferentes enfoques (químico, comportamental o evolutivo). Así, hemos descrito una metodología analítica alternativa para el estudio de las secreciones químicas de lagartos basada en el uso de cromatografía de gases acoplada a espectrometría de masas (GC-MS). Además, hemos estudiado un caso de dimorfismo sexual en la señalización química del lagarto Liolaemus wiegmanni. También investigamos si restricciones en la dieta pueden obstaculizar la expresión de una señal potencialmente honesta (vitamina E) en Iberolacerta cyreni. Además, evaluamos divergencias en la señalización química entre especies relacionadas (Podarcis bocagei y Podarcis carbonelli) y entre poblaciones de la misma especie (Gallotia galloti), discutiendo si el ambiente podría ser el causante de tales diferencias. Finalmente, estudiamos la diversificación de las glándulas foliculares epidérmicas y algunos de los compuestos identificados en las secreciones glandulares a lo largo de Squamata y lacértidos, respectivamente. Así, hemos encontrado que la metodología analítica propuesta es capaz de detectar efectos fisiológicos y ambientales (temperatura) en la composición de las señales química de los lagartos. Por otro lado, las hembras de L. wiegmanni pueden transmitir información a través de las glándulas foliculares y además, esta información es significativamente distinta a la de los machos. Revelamos también que la suplementación en la dieta de vitamina E a los machos de I. cyreni alteró la señalización química y la respuesta inmune de los machos, así como la respuesta comportamental de las hembras. Además, encontramos considerables diferencias en las señales químicas de P. bocagei y P. carbonelli, lo que podría estar asociado a diferencias ambientales entre ambas especies. También, observamos divergencias en el perfil químico de las secreciones femorales de tres poblaciones de G. gallotia (G. g. eisentrauti, G. g. galloti and G. g. palmae), las cuales podrían estar provocadas por diferencias en las condiciones de temperatura y precipitación donde habita cada población. Además, observamos posibles efectos estacionales en los perfiles químicos de G. g. eisentrauti y G. g. galloti. Nuestros análisis sobre la evolución de los ocho compuestos revelaron una diversificación heterogénea de los mismos en cuanto a modo y tiempo. Respecto a las glándulas foliculares epidérmicas en Squamata, nuestros resultados sugieren que el número de estas sigue un modelo de evolución estabilizante, en el que habría un efecto moderado de la señal filogenética. Obtuvimos que la ausencia de estas glándulas sería el estado ancestral en Squamata con un considerable efecto de la filogenia en la localización anatómica de las mismas. El sexo, las condiciones climáticas, la dieta y las relaciones filogenéticas se revelan como factores influyentes en la composición final de las señales químicas de los lagartos y por tanto, el estudio desde múltiples perspectivas es la única vía para conseguir finalmente un profundo conocimiento del papel que juega el sistema quimiosensorial en la ecología y evolución de los lagartos.

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Chemical signals play an important role in intraspecific communication and social organization of many animals, but they also may be useful in interspecific recognition. In lizards, chemical signals are often contained in femoral gland secretions, of which composition may vary between species and populations. This may be especially important in recognition and reproductive isolation between closely related species. We analyzed by gas chromatography-mass spectrometry (GCeMS) the lipophilic fraction of femoral gland secretions of two closely related wall lizard species, Podarcis bocagei and Podarcis carbonelli to test for possible interspecific differences in chemical composition. We found 56 lipophilic compounds in femoral gland secretions of male P. bocagei and 60 in P. carbonelli. The main compounds were steroids and waxy esters, but we also found carboxylic acids and their esters, alcohols, amydes, aldehydes, squalene, ketones and furanones. There were significant differences between species with respect to the number and relative proportions of compounds. Differences in chemical composition might be a consequence of phylogenetic differences per se, but they could also be explained by ecological adaptation to different microclimatic conditions. These differences in chemical profiles may explain the known chemosensory interspecific recognition between these two lizards, contributing to their reproductive isolation.

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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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Records for the amphibian and reptile species of continental Portugal were plotted on maps using the 10 10 km squares of the UTM grid system. A total of 9394 observations were collected, from which 6485 re- cords came from previous national atlases, 1790 from recent published and unpublished work, and 1119 co- rrespond to new observations made by the authors and their collaborators. Overall 31.0% of the records are pre- sented for the first time in a national atlas. Remarkable new records include those for Tarentola mauritanica and Blanus cinereus, enlarging their known range into areas where they were not thought to occur. We present and discuss amphibian and reptile species-density maps.

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Understanding if morphological differences between organisms that occupy different environments are associated to differences in functional performance can suggest a functional link between environmental and morphological variation. In this study we examined three components of the ecomorphological paradigm e morphology, locomotor performance and habitat use e using two syntopic wall lizards endemic to the Iberian Peninsula as a case study to establish whether morphological variation is associated with habitat use and determine the potential relevance of locomotor performance for such an association. Differences in habitat use between both lizards matched patterns of morphological variation. Indeed, individuals of Podarcis guadarramae lusitanicus, which are more flattened, used more rocky environments, whereas Podarcis bocagei, which have higher heads, used more vegetation than rocks. These patterns translated into a significant association between morphology and habitat use. Nevertheless, the two species were only differentiated in some of the functional traits quantified, and locomotor performance did not exhibit an association with morphological traits. Our results suggest that the link between morphology and habitat use is mediated by refuge use, rather than locomotor performance, in this system, and advise caution when extrapolating morphology-performance-environment associations across organisms.

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If morphological variation is tightly related to habitat use, then differences in morphology should affect fitness through their effect on performance within specific habitats. In this study, we investigated intraspecific ecomorphological variation in the generalist lacertid Podarcis bocagei by examining populations exploiting two different habitats: agricultural walls vs. sand dunes, which reflect saxicolous vs. ground-dwelling habits. In the laboratory, we recorded morphological traits, potentially relevant for locomotion and bite performance. We quantified locomotor performance by measuring sprint speed, climbing capacity, maneuverability, and bite force. Based on these measurements, we used univariate and multivariate statistics to examine whether lizards inhabiting the two habitat types differed in morphology and whole-organism performance, and also investigate the association between these two blocks of traits. Our results indicated that lizards from walls and dunes differed in total body size, as well as relative head height. We found significant differences in climbing speed across habitats, where the individuals from walls were better climbers. Finally, bite force was significantly different between habitats where individuals from dunes bite harder than those from walls. A significant association existed between (head) morphology and bite force, but surprisingly, we did not find a significant association between (limb) morphology and locomotor performance. Our results suggest that intraspecific studies are helpful to detect microevolutionary changes.

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A central issue in evolutionary biology is how morphology, performance, and habitat use coevolve. If morphological variation is tightly associated with habitat use, then differences in morphology should affect fitness through their effect on performance within specific habitats. In this study, we investigate how evolutionary forces mold morphological traits and performance differently given the surrounding environment, at the intraspecific level. For this purpose, we selected populations of the lizard Podarcis bocagei from two different habitat types, agricultural walls and dunes, which we expected to reflect saxicolous vs ground-dwelling habits. In the laboratory, we recorded morphological traits as well as performance traits by measuring sprint speed, climbing capacity, maneuverability, and bite force. Our results revealed fast-evolving ecomorphological variation among populations of P. bocagei, where a direct association existed between head morphology and bite performance. However, we could not establish links between limb morphology and locomotor performance at the individual level. Lizards from walls were better climbers than those from dunes, suggesting a very fast evolutionary response. Interestingly, a significant interaction between habitat and sex was detected in climbing performance. In addition, lizards from dunes bit harder than those from walls, although sexual differentiation was definitely the main factor driving variation in head functional morphology. Taking into account all the results, we found a complex interaction between natural and sexual selection on whole-organism performance, which are, in some cases, reflected in morphological variation.

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Biting performance is important for feeding, territory defense, and mating in many animals. While maximal bite force is a well-studied trait, other aspects of biting and their variatio depending on behavior are rarely considered. Here, we took an innovative approach, where (1) we quantified a novel trait, bite duration in lizards; (2) we examined variation across setups that simulate feeding and antagonistic behavior; and (3) we used F-matrix statistics to investigate how different functional components are optimized for ecological and social demands with respect to individual morphology. Our results did not show differences between the sexes in bite duration, but bite performance varied across experimental setups in males, suggesting a higher functional flexibility in this sex. The investigation of form-function associations revealed that trade-offs, facilitations, and one-to-one relationships are simultaneously involved in the morphological optimization of bite force and duration. Put together, our integrated analysis of two different components of bite performance—force and duration—demonstrates their importance for males in both ecological and social tasks. Our findings also suggest the existence of trade-offs in the morphological optimization of functional components, possibly due to physiological constraints on muscle composition, insertion, and orientation.

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The Iberian Peninsula was one of the most important Pleistocene glacial refugia in Europe. A number of recent studies have documented the phylogeography of Iberian taxa and their relationship to more widely distributed species that expanded from this southern European refugium. We use a comparative approach to review the literature that challenges the paradigm of Iberia as a single refuge during Pleistocene glacial maxima and instead supports the occurrence of several Iberian refugia for a range of flora and fauna. Some patterns of phylogeographic concordance were found between the refugial areas identified by different case studies and these broadly overlapped with previously recognized areas of high endemism in the Iberian Peninsula. Such patterns help to illustrate the internal complexity of the Iberian Peninsula as a glacial refugium, and show that for many species, populations with a high degree of genetic structure have existed throughout the Pleistocene. Importantly, the occurrence of these ‘refugia within-refugia’ may confound the interpretation of phylogeographic patterns of European species, and can misleadingly support the occurrence of northern refugia. We discuss these and other consequences, especially when a limited number of samples from the southern European refugia are used.

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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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Wall lizards (Podarcis) are the dominant reptile group across most of southern Europe. Their taxonomy is complex because most species exhibit substantial intraspecific morphological polymorphisms. We have estimated the phylogeny of the particularly diverse western Iberian forms using partial cytochrome oxidase and cytochrome b mitochondrial DNA sequence data and have compared this against morphological variation. Of the two currently recognized species in the area-Podarcis hispanica and P. bocagei-neither is monophyletic, and extremely high genetic diversity between newly identified forms (up to 15% cytochrome b divergences) indicates that both are species complexes. Podarcis b. bocagei is genetically distinct from P. (b.) carbonelli which appears to be a separate species using both mtDNA and protein electrophoretic data. The insular form previously assigned to P. b. berlengensis, and sometimes argued to deserve species status is not genetically distinct from P. (b.) carbonelli using the mtDNA sequences. P. hispanica can be separated into at least four highly divergent groups, two in western Iberia, one in eastern Iberia and one in North Africa.

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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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Several recent papers have reviewed the life and work of French herpetologist Louis Amédée Lantz. They have detailed the composition of his collections deposited in several museums. However, since then, several other important specimens from his collections deposited at the Natural History Museum (NHM, UK) have come to light and we here present all of them in detail. We discovered paralectotypes of Lacerta saxicola obscura Lantz & Cyrén (BMNH 1918.11.21.5–7), syntypes of Lacerta boemica Sukhov (BMNH 1960.1.4.26–30, BMNH 1965.337–342) and Lacerta viridis media Lantz & Cyrén (BMNH 1960.1.4.25, 1966.512). We also identified numerous specimens from the French Mediterranean islands in Lantz’s collection deposited at the NHM, some of which represent the first reported specimens of their species from certain islands. We here provide data on all these specimens. We also place the Mediterranean island specimens from the Lantz collection in their historical context and emphasize the historical and taxonomic value of these collections.

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Understanding the relationship between form and function is central to our comprehension of how phenotypic diversity evolves. Traits involved in multiple activities, such as social interactions and ecological resource use, are under the influence of different evolutionary forces potentially acting in opposite directions. Such systems provide the opportunity of understanding how potential constraints on morphological variation may influence whole-organism performance. In this study we examined morphology and bite performance in two closely related species of Podarcis wall lizards with divergent microhabitat preferences, to investigate how natural and sexual selection interact to shape the evolution of head traits. Our results show that although head morphology is markedly different between species and sexes, only sexes differ in bite force, indicating that the ecological differentiation between species is reflected in their morphology but does not constrain performance. Rather, the modification of the relative size of head components between species and a shift in the form-function relationship provide a potential explanation of how equal performance is attained by different morphological configurations. Geometric morphometrics provide a clear, biomechanically meaningful image of how this is achieved and show a bisexual pattern of head shape-bite force association in both species. This, together with a strong allometry of head size on body size and head shape on head size, provides indirect morphological evidence for the importance of sexual selection in shaping morphological and functional patterns. Finally, our findings suggest that the differences observed between species and sexes in head traits and bite performance are not reflected in their dietary ecology, implying that if trophic niche segregation between groups occurs, the reasons behind it are not primarily related to head morphology and functional variation.

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Sexual dimorphism (SD) is the result of evolutionary pressures acting differentially on members of each sex. Investigating the association between SD observed in different sets of phenotypic traits, which are evolutionarily linked, can shed light on the mechanisms causing SD variation within and across species. Although the association between morphology and locomotor performance is a major paradigm in ecomorphology, substantially less effort has been dedicated to investigate the covariation between both sets of traits in the context of sexual divergence. Here, we investigated morphology and locomotor performance in wall lizards Podarcis bocagei to determine if locomotor SD exists in this species, as one may expect based on the morphological SD observed, and test whether both types of SD are directly associated. Our results indicate that significant morphological and locomotor SD exists in this species, reporting a significant locomotor SD for the first time in this genus of lizards. Our study also provides evidence that a direct association between morphology and performance exists at the individual level, binding together SD in both sets of traits. The observed patterns of SD suggest that male locomotor capacity and the corresponding morphological features are well suited for sprinting in level surfaces, but less so for other types of locomotion, potentially as a result of sexual selection acting on male locomotor performance through influences on territory defence and reproductive fitness.

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Sexual dimorphism (SD) in locomotor performance is prominent in many species of lizards. This functional pattern is related to both size and shape SD and it has been associated to sexual selection on males, related to territorial behaviour. Locomotor capacities of lacertid lizards are related to habitat variation, following ecomorphological predictions, but SD has not been extensively investigated. The few existing studies have indicated a lack of SD in this respect. We investigated SD in locomotor performance and its relation to morphological traits in a population of Podarcis bocagei, a species endemic to the Iberian Peninsula. We measured sprinting, climbing and clambering speed, as well as manoeuvrability, of adult male and female P. bocagei in a racing track. We also quantiied body size and shape, including characters potentially relevant for locomotion, in the same individuals. Our results indicate that while SD in performance is not signiicant when looking at each locomotor component separately, diferences between sexes are signiicant when treating the data in a multivariate framework. This is due to diferences between sexes in the trade-ofs between conditions: while females are relatively stable across designs, males are fast sprinters, but sufer a visible reduction of their locomotor capacity when climbing or serpentizing. Fitting biomechanical predictions and previous observations, the morphological variables associated to locomotor performance are total body size and limb length. Our study suggests that signiicant, but previously unnoticed, SD may exist in locomotor performance in lacertids and remarks the importance of multivariate analyses when examining performance capacities.

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Morphological variation in relation to habitat is known to occur in several lizard groups. Comparative studies have linked morphology and habitat use, showing that locomotion is the principal mediator of this evolutionary relationship. We investigated intraspecific ecomorphological variation in Podarcis bocagei by examining three habitat types, representing a variety between saxicolous and ground-dwelling habits. Our results indicate variation in absolute and relative limb length, but patterns are only partially concordant to biomechanical predictions. While the femur and hind foot are longer in ground-dwelling lizards, confirming previous observations, the tibia and hind limb are relatively shorter, contradicting expectations. Additionally, head shape varies substantially between habitats, in line with a hypothesis of mechanical restrictions related to microhabitat and refuge use. Finally, we detect male-specific variation between habitats in total body size and head size, providing evidence for interactions between natural and sexual selection. Interestingly, the results obtained about head size and shape partially coincide with the interspecific ecomorphological variation across the whole genus Podarcis. Although performance and behaviour studies are necessary to definitely confirm the functional and evolutionary significance of the observed patterns, our study indicates that ecomorphological adaptations can arise in a very short evolutionary time in this group of lizards.

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Podarcis bocagei and Podarcis carbonelli are two closely related species, endemic to the western Iberian Peninsula, whose biometric differences have already been established. However, their pholidosis has not been investigated in detail. We studied various pholidotic characters, often used for taxonomic purposes in the Lacertidae, in order to analyse scalation patterns in the two species and detect characters that could be useful for their diagnosis. The results show that both species present significant differences in various pholidotic characters, although patterns differ between the sexes. The combination of continuous and meristic characters allowed a good discrimination between P. bocagei and P. carbonelli, especially when sexes were separated a priori. However, the close phylogenetic relationship between the two species, along with their high intraspecific morphological variability, complicate the use of pholidotic characters as diagnostic tools in the field.

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Podarcis bocagei and P. carbonelli are two closely related lacertid species, very similar morphologically and ecologically. We investigated sexual dimorphism patterns presented by both species in allopatry and in sympatry. Sexual size and shape dimorphism patterns were analyzed using both multivariate and geometric morphometric techniques. Multivariate morphometrics revealed a marked sexual dimorphism in both species--males being larger with more robust habitus and females presenting a longer trunk. General patterns of sexual size dimorphism are not modified in sympatry, although there is evidence for some morphological change in male head size. The application of geometric morphometrics offered a more detailed image of head shape and revealed that males present a more developed tympanic area than do females, while females have a more rounded head. Differences in the degree of sexual shape dimorphism were detected in sympatry, but no consistent patterns were observed. From the results of the study, and based on previous knowledge on the populations studied, we conclude that the morphological differences observed are probably not caused by exploitative competition between the species, but rather appear attributable to the modification of the relative influence of sexual and natural selection on both sexes.

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Podarcis bocagei and Podarcis carbonelli are two lacertid species endemic to the western Iberian Peninsula, and both show head size and shape sexual dimorphism. We studied immature and adult head sexual dimorphism and analysed ontogenetic trajectories of head traits with body and head size, aiming to shed light on the proximate mechanisms involved. Immatures were much less dimorphic than adults, but geometric morphometric techniques revealed that head shape sexual differences are already present at this stage. Males and females differed in allometry of all head characters with body size, with males showing a disproportionate increase of head size and dimensions. On the other hand, head dimensions and head shape changed with increasing head size following similar trends in both sexes, possibly indicating developmental restrictions. Consequently, adult sexual dimor- phism for head characters in these species is the result of both shape differences in the immature stage and hypermetric growth of the head in relation to body size in males.

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Podarcis bocagei and P. carbonelli are two closely related lizard species, endemic to the western Iberian Peninsula. Although genetic and morphological distinction between them is now well established, information on anatomical features is still very restricted. We studied presacral vertebrae number in both species, separately examining cervical and trunk vertebrae counts, in order to assess the interspecific and intersexual variation in these characters. There was no significant interspecific variation among lizards of the same sex for any of the vertebrae counts studied. However, important sexual variation existed, females presenting higher counts than males for both trunk and total presacral vertebrae, while males showed higher values of cervical vertebrae. Although our results lie within the known limits for other closely related species, they reveal that vertebrae number might present higher intraspecific variation than previously thought.

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Sexual dimorphism in body size and shape in animals is normally linked to sexual selection mechanisms that modify the morphological properties of each sex. However, sexual dimorphism of ecologically relevant traits may be amplified by natural selection and result in the ecological segregation of both sexes. In the present study, we investigated patterns of sexual dimorphism of morphological traits relevant for locomotion in two lacertid lizards, Podarcis bocagei and Podarcis carbonelli, aiming to identify ontogenetic sources of variation. We analysed trunk and limb variation in relation to total body size, as well as the covariation of different traits, aiming to shed light on the proximate causation of adult sexual dimorphism. We find that, although immatures are generally monomorphic, adult females have a longer trunk, and adult males have longer fore and hind limbs. Both sexes differ substantially with respect to their growth trajectories and relationships between traits, whereas, in some cases, there are signs of morphological constraints delimiting the observed patterns. Because of the direct connection between limb size/shape and locomotor performance, which is relevant both for habitat use and escape from predators, the observed patterns of sexual dimorphism are expected to translate into ecological differences between both sexes.

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Morphological variation in relation to habitat is known to occur in several lizard groups. Comparative studies have linked morphology and habitat use, showing that locomotion is the principal mediator of this evolutionary relationship. Here, we investigate intraspecific ecomorphological variation in Podarcis bocagei by examining three habitat types, representing a variety between saxicolous and ground-dwelling habits. Our results indicate variation in absolute and relative limb length, but patterns are only partially concordant to biomechanical predictions. Whereas the femur and hind foot are longer in ground-dwelling lizards, confirming previous observations, the tibia and hind limb are relatively shorter, contradicting expectations. Additionally, head shape varies substantially between habitats, in line with a hypothesis of mechanical restrictions related to microhabitat and refuge use. Finally, we detect male-specific variation between habitats in total body size and head size, providing evidence for interactions between natural and sexual selection. Although performance and behaviour studies are necessary to definitely confirm the functional and evolutionary significance of the observed patterns, our study indicates that ecomorphological adaptations can arise in a very short evolutionary time in this group of lizards.

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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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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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Background: Understanding how phenotypic variation scales from individuals, through populations, up to species, and how it relates to genetic and environmental factors, is essential for deciphering the evolutionary mechanisms that drive biodiversity. We used two species of Podarcis wall lizards to test whether phenotypic diversity within and divergence across populations follow concordant patterns, and to examine how phenotypic variation responds to genetic and environmental variability across different hierarchical levels of biological organization, in an explicit geographic framework. Results: We found a general concordance of phenotypic variation across hierarchical levels (i.e. individuals and populations). However, we also found that within-population diversity does not exhibit a coherent geographic structure for most traits, while among-population divergence does, suggesting that different mechanisms may underlie the generation of diversity at these two levels. Furthermore, the association of phenotypic variation with genetic and environmental factors varied extensively between hierarchical levels and across traits, hampering the identification of simple rules to explain what yields diversity. Conclusions: Our results in some cases comply with general ecological and evolutionary predictions, but in others they are difficult to explain in the geographic framework used, suggesting that habitat characteristics and other regulatory mechanisms may have a more substantial contribution in shaping phenotypic diversity.

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Among the Iberian members of the genus Podarcis, two species are present in the North-Western region: P. bocagei and P. carbonelli. The high morphological and ecological similarity between them as well as their well-known distribution ranges, provides a special opportunity to evaluate asymmetry patterns typically displayed by wall-lizards. Levels of asymmetry expression have been correlated with several kinds of abiotic, biotic and genetic stresses. The few asymmetry studies focusing on reptiles have found that even minor morphological asymmetries could have an impact on individuals’ survival, altering their fitness for example through reduced escaping performance. Others found that asymmetry varied across the species distribution range, increasing in the peripheral populations, while others evaluated isolation effects between insular and continental populations through fluctuating asymmetry. Therefore, asymmetry may be a useful fitness index but in order to be valuable in comparative studies, the levels of variation usually displayed by natural populations must be determined first. In our study we aim to define the asymmetry patterns present in these species through the analysis of meristic and linear biometric bilateral traits. We include characters from different structures, like the head and the limbs, to capture individual asymmetric variation. Measurement error was assessed in order to quantify the between sides variation truly due to asymmetry. After this initial stage, those traits that showed fluctuating asymmetry were analyzed separately because these can be used to evaluate the levels of developmental instability displayed by different populations. The occurrence of other types of asymmetry, particularly directional asymmetry, were also detected and evaluated. The similarity between asymmetry patterns of both species was tested as well as the occurrence of sexual dimorphism in asymmetry variation. We exemplify the application of asymmetry indices to a specific situation of sympatry, as it may pose stressful conditions to populations occupying similar habitats. The results will be presented in the ecological and evolutionary framework of these species.

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Iberian Podarcis have been the target of intensive phylogenetic research during recent years because of the previously unexpected diversity found within. At present, the former Podarcis hispanica is recognized as a species complex parafiletic to P. bocagei, P. carbonelli and P. vaucheri with the majority of lineages being strongly supported by evidence coming from mtDNA, nuclear DNA and protein electrophoretic data. In some cases, morphological studies focusing on specific lineages have also agreed with their phenotypic distinctiveness which consequently conducted to their raise to the specific rank, such as P. carbonelli and P. vaucheri. However, some of the recent identified forms (i.e. “morphotype 1”) still contain considerable genetic diversity, while the apparent relationships of other monophyletic groups change according to the kind of data analyzed. From the recently published data, one highly divergent lineage seems to deserve a deeper look: Podarcis “morphotype 2”. This lineage is the most widespread in the Iberian Peninsula, reported from across the Central and Southwestern regions. While its distribution in Portugal is well defined, its eastern distribution is less well known. Reflecting this situation, recent studies lack representative samples from its entire distribution range. However, when mtDNA markers were preliminary analyzed, considerable variation was reported but without obvious geographic partitioning. With this study we aim to evaluate the genetic diversity within this form. We conducted a more complete sampling scheme trying to cover the entire distribution range of this specific lineage. We combine new and previously published sequence data in order to perform a more complete phylogeographic analysis of the whole lineage. Furthermore, the geographic distribution of this form is now more precisely delimited.

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Podarcis bocagei y Podarcis carbonelli son dos especies cercanas de lacértidos endémicos del oeste de la Península Ibérica, cuyas diferencias biométricas ya han sido demostradas. Sin embargo, su folidosis nunca se ha estudiado detalladamente. Hemos estudiado diversos caracteres folidóticos, habitualmente utilizados en la taxonomía de lacértidos, con el objetivo de analizar los patrones que caracterizan ambas especies y detectar aquellos que podrían emplearse para su distinción. Los resultados demuestran que ambas especies presentan diferencias significativas en varios caracteres, aunque los patrones observados varían entre los sexos. La combinación de caracteres cuantitativos y cualitativos permitió una aceptable discriminación entre P. bocagei y P. carbonelli, especialmente cuando los sexos se separaron a priori. Sin embargo, las estrechas relaciones filogenéticas entre ambas especies y su elevada variabilidad morfológica intraespecífica dificultan la selección de caracteres que podrían servir como herramientas de diagnosis en el campo.

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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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Regeneration is the process of regrowth of an injured/missing tissue/organ from the residual tissue. One outstanding example is the regeneration of tail in lizards in which they can completely regenerate without the formation of scaring or fibrotic tissue. This study presents an analysis of the main differences between a mature regenerated tail and the original tail of the lizard Podarcis bocagei by comparing histological biomarkers for support structures, morphology and pigmentation, vasculature and energy storage. Matured regenerated and original tails were studied with different histological staining’s and immunohistochemistry to highlight the various tissue components. We found differences in the morphological features scalation pattern and pigmentation, as well as in the support structures: muscle organization, collagen and calcification. Vasculature was remodeled. Energy storage was restored with regeneration maintaining therefore the main metabolic function. We demonstrate that the regenerated tail does not lose its main functions like locomotion and energy metabolism. The present findings may open a new window of research with implications in regenerative medicine.

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Identifying factors influencing infection patterns among hosts is critical for our understanding of the evolution and impact of parasitism in natural populations. However, the correct estimation of infection parameters depends on the performance of detection and quantification methods. In this study, we designed a quantitative PCR (qPCR) assay targeting the 18 S rRNA gene to estimate prevalence and intensity of Hepatozoon infection and compared its performance with microscopy and PCR. Using qPCR, we also compared various protocols that differ in the biological source and the extraction methods. Our results show that the qPCR approach on DNA extracted from blood samples, regardless of the extraction protocol, provided the most sensitive estimates of Hepatozoon infection parameters; while allowed us to differentiate between mixed infections of Adeleorinid (Hepatozoon) and Eimeriorinid (Schellackia and Lankesterella), based on the analysis of melting curves. We also show that tissue and saline methods can be used as low-cost alternatives in parasitological studies. The next step was to test our qPCR assay in a biological context, and for this purpose we investigated infection patterns between two sympatric lacertid species, which are naturally infected with apicomplexan hemoparasites, such as the genera Schellackia (Eimeriorina) and Hepatozoon (Adeleorina). From a biological standpoint, we found a positive correlation between Hepatozoon intensity of infection and host body size within each host species, being significantly higher in males, and higher in the smaller sized host species. These variations can be associated with a number of host intrinsic factors, like hormonal and immunological traits, that require further investigation. Our findings are relevant as they pinpoint the importance of accounting for methodological issues to better estimate infection in parasitological studies, and illustrate how between-host factors can influence parasite distributions in sympatric natural populations.

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The genus Hepatozoon Miller, 1908 (Apicomplexa: Adeleorina) is composed of intracellular haemogregarine parasites that are widely distributed among all tetrapod groups. The present study combines microscopic and molecular data on haemogregarine parasites from lizards in the western Mediterranean. We screened tissue samples and examined blood smears for the presence of species of Hepatozoon from four lizards, namely Algyroides marchi Valverde, endemic to Southeast Spain, Podarcis bocagei Seoane from Spain and Portugal, P hispanica Steindachner from Spain, and P lilfordi Günther from Cabrera, Balearic Islands (Spain). Our results show that prevalence and intensity of Hepatozoon parasites vary between and within lizard species from different regions. Algyroides marchi and P bocagei from Spain had the lowest values, whereas P hispanica had the highest. Phylogeny based on 18S rRNA gene sequences indicates that most of the new Hepatozoon sequences are part of a clade exclusive from North African and Iberian lizards, except for a single P bocagei isolate that is found related to another clade including isolates from other reptile host species and rodents. Interestingly, isolates from Algyroides form a distinct monophyletic subgroup, which could be a signal of strict host-specificity within this host genus.

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This survey is presenting the geographical range of 17 amphibian and 29 reptile species (including the marine turtles) known to occur in Portugal, based on available data from the literature as well as ehe author`s own observations during the years 1976- 1981. It is the first attempt to map exactly the herpetofauna of a southern European country. Data are given concerning the horizontal and vertical distribution of each species, the density of distribution as well as the typical habitats, the latter being described in some detail. The rhythm of activity of most of the species depends on the different climatic conditions within this country. The few available phenological data are sum- marized.

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The results of two excursions (winter 1982/83 and spring 1983) to Portugal require the present additional note to my paper: `Contribution to the knowledge of the distribution of the Portuguese herpetofauna.` Remarkable new distributional data are provided for Chioglossa lusitanica, Alytes cisternasii, Pelobates cultripes, Tarentola mauritanica, Podarcis hispanica and Psammodromus hispanicus.

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Zwischen 1976 und 2002 wurde die Artenzusammensetzung der Herpetofauna an 90 Ruinen von Wassermühlen in Portugal untersucht. Es wurden 12 Amphibien- und 19 Reptilienarten registriert, die 69 % der in Kontinental-Portugal vorkommenden Arten repräsentieren. Die Zusammensetzung der Herpetozönosen wird durch die geografische Lage, den Habitattypus des Umfeldes und der Phase des Zerfallsprozesses der einzelnen Mühlen geprägt. Durch die anthropogene Belastung der Flüsse ist vielerorts eine deutliche Reduzierung der Individuendichte und Artenzahl der Amphibien und hydrophiler Reptilien zu verzeichnen.

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Portugal ist ein beliebtes Urlaubsland. Vor allem die Metropole Lissabon und die Atlantikstrände des Algarve ziehen Touristen an. Aber das Land an der Südwestspitze Europas hat noch viel mehr zu bieten. Die Serra de Montemuro ist ein lohnendes Reiseziel auch für alle, die nach Amphibien und Reptiulien Ausschau halten.

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Wir stellen die Verbreitungsverhältnisse der Amphibien und Reptilien der Barroso-Region (Nordportugal) vor. Einleitend werden die geologischen und klimatischen Bedingungen des Untersuchungsgebietes beschrieben, die Habitattypen und die sich auf den Kartierungsraum beziehende herpetofaunistische Literatur vorgestellt. Bisher wurden 12 Amphibien- und 19 Reptilienarten registriert. Ihre Verbreitung wird kartografisch auf der Basis eines 1 x 1 km UTM-Gitternetzes dargestellt. Charakteristisch für den gesamten Untersuchungsraum ist die Dominanz eurosibirischer Arten mit zahlreichen nordiberischen Endemiten (Chioglossa lusitanica, Rana iberica, Lacerta schreiberi, Podarcis bocagei, Vipera seoanei), unter die sich an der klimabegünstigten Südabdachung einige mediterrane Arten (Tarentola mauritanica, Blanus cinereus) mischen. Vipera seoanei erreicht im Untersuchungsgebiet das am weitesten nach Süden vorgeschobene Vorkommen ihres Gesamtareals; Psammodromus hispanicus und Chalcides bedriagai die nordwestliche Grenzzone ihrer Verbreitung in Portugal. Abschließend wird auf die Bedeutung des Einflusses der anthropogenen Landschaftsgestaltung auf die Verbreitungsmuster der einzelnen Arten hingewiesen und die aktuellen Gefährdungsursachen (Landnutzungswandel, Wiederbewaldung, Talsperrenbau) beschrieben.

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We describe the distribution of the amphibians and reptiles of the Serra de Monte-muro (northern Portugal). In the introduction the geological and climatical condi-tions of this region as well as the habitat types and the herpetofaunistic studies are described. 11 amphibian and 17 reptile species were recorded. The distribution pat-tern of each species is shown by maps using the 1 x 1 km squares of the UTM grid system. In the montane region predominate north Iberian endemits (Chioglossa lusi-tanica, Rana iberica, Lacerta schreiberi; the occurrence of Podarcis bocagei is doubtful) and ubiquists (Salamandra salamandra, Triturus marmoratus, Lissotriton boscai, Pelophy-lax perezi; Chalcides striatus, Timon lepidus, Podarcis guadarramae). Along the rivers (Rio Douro, Rio Paiva), areas with components of Mediterranean climate, thermophile species (Chalcides bedriagai, Blanus cinereus, Hemorrhois hippocrepis) penetrate into the investigated area. Within the region Podarcis carbonelli reaches the northernmost oc-currence of its total distribution range. Remarkable is the high number of individuals of some montane Hyla molleri populations. Finally, the human impact on the land-scape (change of agricultural activities, reforestation in monocultures, widespread forest fires), its influence on the distribution pattern of the amphibians and reptiles and the efforts for protect the habitats are discussed.

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1. Female and male hosts may maximise their fitness by evolving different strategies to compensate for the costs of parasite infections. The resulting sexual dimorphism might be apparent in differential relationships between parasite load and body condition, potentially reflecting differences in energy allocation to anti-parasitic defences. For example, male lacertids with high body condition may produce many offspring while being intensely parasitised. In contrast, female lacertids may show a different outcome of the trade-offs between body condition and immunity, aiming to better protect themselves from the harm of parasites. - 2. We predicted that females would have fewer parasites than males and a lower body condition across parasitaemia levels because they would invest resources in parasite defence to mitigate the costs of infection. In contrast, the male strategy to maximise access to females would imply some level of parasite tolerance and, thus, higher parasitaemia. - 3. We analysed the relationship between the body condition of lizards and the parasitemias of Karyolysus and Schellackia, two genera of blood parasites with different phylogenetic origins, in 565 females and 899 males belonging to 10 species of the Lacertidae (Squamata). These lizards were sampled over a period of 12?years across 34 sampling sites in southwestern Europe. - 4. The results concerning the Karyolysus infections were consistent with the predictions, with males having similar body condition across parasitaemia levels even though they had higher infection intensities than females. On the other hand, females with higher levels of Karyolysus parasitaemia had lower body condition. This is consistent with the prediction that different life strategies of male and female lacertids can explain the infection patterns of Karyolysus. In contrast, the parasitaemia of Schellackia was consistently low in both male and female hosts, with no significant effect on the body condition of lizards. This suggests that lizards of both sexes maintain this parasite below a pathogenic threshold.

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Report on two excursions to the Iberian Peninsula in 1981 and 1982: Pyrenees, Cantabrian Mountains, Galicia, Peneda-Geres (Portugal), Chaves (Northeast of Portugal), Monfragüe (Estremadura), Andalusia, East-Algarve, Toledo and Argeles sur Mer (South-France). A total of 37 species of amphibians and reptiles could be observed (urodelans 6, anurans 8, chelonians 1, lizards 14 and snakes 8). The visited areas are briefly characterized in terms of their herpetocoenosis. Information on habitat requirements, partly also on body measurements of the species recorded are given. New data on the range limits of Lacerta schreiberei, L. lepida, L. vivipara, L. monticola and Psammodromus algirus are given. A syntopic occurrence of Coronella austriaca and C. girondica in the Cantabrian Mountains was found. C. girondica is recorded for the first time in Asturia. A remarkable finding in the Cantabrian Mountains was a melanistic Aesculapian Snake (Elaphe longissima). This is the second record of this species in Spain. In these Northern Spanish highlands the breeding season of the amphibians is extended into summer. In North Portugal we observed Natrix maura eating Chioglossa lusitanica.

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Aim Body size is instrumental in influencing animal physiology, morphology, ecology and evolution, as well as extinction risk. I examine several hypotheses regarding the influence of body size on lizard evolution and extinction risk, assessing whether body size influences, or is influenced by, species richness, herbivory, island dwelling and extinction risk. Location World-wide. Methods I used literature data and measurements of museum and live specimens to estimate lizard body size distributions. Results I obtained body size data for 99% of the world`s lizard species. The body size–frequency distribution is highly modal and right skewed and similar distributions characterize most lizard families and lizard assemblages across biogeographical realms. There is a strong negative correlation between mean body size within families and species richness. Herbivorous lizards are larger than omnivorous and carnivorous ones, and aquatic lizards are larger than non-aquatic species. Diurnal activity is associated with small body size. Insular lizards tend towards both extremes of the size spectrum. Extinction risk increases with body size of species for which risk has been assessed. Main conclusions Small size seems to promote fast diversification of disparate body plans. The absence of mammalian predators allows insular lizards to attain larger body sizes by means of release from predation and allows them to evolve into the top predator niche. Island living also promotes a high frequency of herbivory, which is also associated with large size. Aquatic and nocturnal lizards probably evolve large size because of thermal constraints. The association between large size and high extinction risk, however, probably reflects a bias in the species in which risk has been studied.

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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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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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Mountain lizards are highly vulnerable to climate change, and the continuous warming of their habitats could be seriously threatening their survival. We aim to compare the thermal ecology and microhabitat selection of a mountain lizard, Iberolacerta galani, and a widely distributed lizard, Podarcis bocagei, in a montane area. Both species are currently in close syntopy in the study area, at 1,400 m above the sea level. We determined the precision, accuracy and effectiveness of thermoregulation, and the thermal quality of habitat for both species. We also compared the selection of thermal microhabitats between both species. Results show that I. galani is a cold-adapted thermal specialist with a preferred temperature range of 27.929.7 C, while P. bocagei would be a thermal generalist, with a broader and higher preferred temperature range (30.134.5 C). In addition, I. galani selects rocky substrates while P. bocagei selects warmer soil and leaf litter substrates. The thermal quality of the habitat is higher for P. bocagei than for I. galani. Finally, P. bocagei achieves a significantly higher effectiveness of thermoregulation (0.87) than I. galani (0.80). Therefore, these mountain habitat conditions seem currently more suitable for performance of thermophilic generalist lizards than for cold-specialist lizards.

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The development of molecular methods is becoming a promising field in the analysis of parasite fauna in wildlife species. This is especially useful in the case of parasite species where developmental larval stages are difficult to assess using standard methods. In this study we screened for the presence of parasitic nematodes infecting lacertid lizards from the Iberian Peninsula and Balearic islands using nematode-specific 18S ribosomal RNA gene primers on host tissue samples. Sequencing of positive samples revealed the presence of different genera of nematodes. The detection of Strongyloides, a monoxenous genus reported for the first time in Podarcis lilfordi is probably the result of the amplification of larval stages present in the host circulatory system. Two spirurid nematodes, Synhimantus and a new unidentified clade, were also found, suggesting that reptiles might be paratenic hosts of several spirurid species. This study demonstrates the benefits of using specific molecular markers on tissue samples to identify infecting larval stages of nematodes, otherwise difficult to assess using traditional screening methods.

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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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P. bocagei is considered a valid species, and several criteria are used for distinguish it in sympatry of P. hispanica, a closely related species. A detailed distribution map is provided for this lizard, which occurs in Galicia, León, Orense, Zamora, western slopes of the Sistema Central, and probably, the north of Portugal. P. hispanica males and females in the Sistema Central are significant larger than males and females of P. bocagei. Significant differences in size, scalation and ecology are apparent between the Sistema Central and northern populations of P. bocagei, but no taxonomic decisions are made, pending a more detailed study. The examination of the masseteric shield revealed its large size in P. bocagei and its absence in most of the P. hispanica studied. P. bocagei has a relatively robust, deep skull, probably related to his terrestriral habits, unlike P. hispanica, which has a flattened skull and, at least in the Sistema Central, is a saxicolous species. P. bocagei is very common in Galicia and is found there at altitudes between 0 and 1500 meters, but in the Sistema Central is restricted to Quercus pyrenaica forests, between 800 and 1200 meters.

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The taconomic status of Podarcis bocagei in the Iberian Peninsula is reviewed. Since no Holotype is known, a Lectotype from the species is given. P. bocagei carbonelli ssp. n. is also described; it inhabits the western area of the Sistema Central, in particular the `Sierras` of Francia and Gata, both in Spain and the Serra da Estrela in Portugal. Finally, new localities for both subspecies are described, that together with those reported in the previous paper sum a total of 76 localities known up to date.

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A study on taxocenosis of Lacertidae with 8 species: Lacerta lepida, Lacerta schreiberi, Lacerta montícola, Acanthodactylus erythrurus, Psammodromus algirus, Psammodromus hispanicus, Podareis hispanica and Podareis bocagei in the western part of the Sistema Central (Iberian Península) has been carried out. Most of these behave as feeding generalists but A. erythrurus seems to specialize in mymecophagy whereas L. montícola turns out to be a «forced» sthenophage. The overlap valúes in each niche dimensión suggest that the spatial dimensión and habitat preference are the most important factors. However, segregation is produced jointly due to the interaction of the three dimensions considered which act interdependently.

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Tail autotomy is one of the main anti-predator mechanisms of lacertid lizards, but it has been predicted that it is only retained in its full capacity when its benefits exceed its costs (Arnold, 1988). To test this hypothesis, ease of tail shedding was examined in a number of continental and insular lacertid lizard populations, each of which showed a different shedding capacity. Tails are shed more easily in those continental and insular populations where there is a greater probability of predation. In insular populations not subjected to strong predation, the tail tends to be retained. The relationship of these findings to insular Mediterranean lizard populations and to the extinction of the Balearic lizard, Podarcis lilfordi are discussed.

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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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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. Conversely, the study of two nuclear gene genealogies (β-fibint7 and 6-Pgdint7), demonstrated that, in general, species and mtDNA lineages are strikingly non- monophyletic, 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 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 by- product 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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Podarcis bocagei and P. carbonelli are two Iberian-endemic wall lizard species for which a detailed phylogeographic study based on mitochondrial DNA (mtDNA) variation has been accomplished, showing that the two species responded differently to climatic oscillations during the Quaternary. In this study, our main goal was to assess whether nuclear markers reveal similar evolutionary patterns to those inferred from the study of mtDNA variation. We studied a battery of allozyme and microsatellite loci in populations from both species. For each species we evaluated overall levels of differentiation, the patterns of variation in genetic variability, genetic relationships among populations, changes in allele frequencies across the species’ distribution area and applied model-based individual multilocus genotype clustering approaches to detect hidden population structure. Our results for Podarcis bocagei are highly concordant with the phylogeographic scenario inferred from mtDNA variation: not only we find very low levels of population differentiation, consistent with survival in a single glacial refugium, but we detect signatures of a rapid demographic and geographic expansion, as suggested from mtDNA analyses. The analyses of nuclear markers furthermore helped in the identification of the probable refugial area, as well of expansion routes. Also with similarity to observations based on mtDNA variation, a low level of population differentiation was observed in P. carbonelli. In concordance with mitochondrial DNA, differentiation levels were slightly higher in P. carbonelli than those observed in P. bocagei. However, the geographic basis for differentiation in P. carbonelli is highly inconsistent between mtDNA and nuclear markers, suggesting a complex, albeit recent, history of isolation. Furthermore, a recent reduction on the species distribution has probably erased the signatures of glacial isolation and postglacial expansion that are normally found in other Iberian species, suggesting that the currently observed pattern of genetic differentiation in this species was shaped more by recent genetic drift than by the Pleistocene climatic oscillations.

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The taxonomy of Iberian and North African wall lizards (Podarcis sp.) has been controversial. Recently, morphological and mtDNA sequence data have provided new information on differentiation within these lacertids. To compare these results to those provided by nuclear markers, we investigated variation at 11 polymorphic protein loci using conventional electrophoresis and isoelectric focusing in 11 populations belonging to seven different mtDNA lineages. A total of 62 alleles were found. Populations belonging to the same mtDNA type presented high genetic similarity, whereas strong differentiation was observed between groups. These results are consistent with those previously obtained from morphological and mtDNA analysis and support the idea that Iberian and North African Podarcis are composed of several well-differentiated entities, some of which are already recognized as species, whereas others (belonging to the P. hispanica complex) clearly need taxonomic revision.

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Pleistocene climatic oscillations were a major force shaping genetic variability in many taxa. We analyse the relative effects of the ice ages across a latitudinal gradient in the Western Mediterranean region, testing two main predictions: (i) species with historical distributions in northern latitudes should have experienced greater loss of suitable habitat, resulting in higher extinction of historical lineages than species distributed in southern latitudes, where the effects of the ice ages were not as drastic. This would be reflected in the observation of lower diversity and number of differentiated lineages in northern areas. (ii) a signature of demographic expansion following the climate amelioration should be obvious in northern species, whereas in the south evidence of long-term effective population size stability should be observed. We used as models three species of wall lizards (Podarcis bocagei, Podarcis carbonelli and Podarcis vaucheri) that replace each other along the study area. We investigated the patterns of mitochondrial DNA diversity and subdivision and obtained demographic parameter estimates for each species. Our results suggest that P. bocagei, the northernmost species, bears low genetic diversity, a shallow coalescent history and marks of a demographic expansion. In contrast, P. vaucheri, the species with a southernmost distribution, shows deeper coalescence events, complex geographical substructure and no evidence for population growth. The species with an intermediate distribution, P. carbonelli, shows average levels of diversity, substructure and population growth. Taken together, these results conform to our main predictions and are explained by a differential influence of the ice ages on distinct latitudes.

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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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When recently diverged taxa come into contact, the extent of introgression between them is related to the degree of differentiation that they have achieved. Studying contact zones is therefore essential to understand if differentiated taxa are reproductively isolated and, ultimately, if they are likely to remain distinct. Recent work on Iberian and North African wall lizards (Podarcis) has documented the existence of multiple evolutionary units, diagnosable both by genetic markers and morphology, but suggests that gene flow between distinct forms has occurred. Therefore, we were interested in evaluating how species boundaries are maintained in the areas where they meet. In this work, we study the contact zone between Podarcis bocagei and Podarcis carbonelli. We sampled a transect including the only locality where these two lizards are known to occur in syntopy and analysed a battery of 15 unlinked nuclear genetic markers and mitochondrial DNA. We also conducted a preliminary analysis of morphology and fertility. Using model-based clustering approaches, we show that the two species hybridize in the population where they have direct contact, but evidences of introgression are low for nearby populations. Although a significant number of individuals show evidence of admixture, this hybrid zone is clearly bimodal, suggesting strong barriers to gene flow, of which the putative nature are discussed. Interestingly, morphological analyses do not support the existence of intermediate forms among individuals that are admixed genetically. Taken together, these results constitute further evidence validating P. bocagei and P. carbonelli as distinct species.

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When recently diverged taxa come into contact, the extent of introgression between them is directly related to the stage of differentiation that they have achieved. The study of contact zones is therefore essential to understand if differentiated taxa are reproductively isolated and, ultimately, to assess if they are likely to remain distinct. Recent work on differentiation within Iberian and North African wall lizards (Podarcis) has shown that multiple evolutionary units exist, diagnosable both by genetic markers and morphology, some of which have been recognised as full species. These studies also suggest that gene flow between distinct forms may occur. Therefore, we were interested in evaluating how species boundaries are maintained in the areas where they meet. In this work, we present data relative to the contact zone between P. bocagei and P. carbonelli. We studied a transect including the only known locality where these two lizards are known to occur in syntopy and analysed a battery of 15 unlinked nuclear genetic markers (11 allozymes, three microsatellites and one nuclear intron), mitochondrial DNA and morphology. We also conducted a preliminary analysis of fertility. Using model-based clustering approaches we show that the two species hybridize in the population where they contact, but evidences of introgression are low for nearby populations. Although a significant number of individuals show evidence of admixture, this hybrid zone is clearly bimodal, suggesting strong barriers to gene flow, which putative nature are discussed. Interestingly, morphological analyses revealed a complete lack of intermediate forms, even among individuals that are admixed genetically. Taken together, these results constitute further evidence validating the species status of P. bocagei and P. carbonelli.

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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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Podarcis bocagei and Podarcis carbonelli are two species of wall lizards endemic to the western Iberian Peninsula. A detailed phylogeographical study based on mitochondrial DNA (mtDNA) variation has shown that they responded differently to the Quaternary climatic oscillations. These differences have been attributed to their distribution patterns: P. bocagei is distributed in the north of the Peninsula and in a continuous fashion, whereas P. carbonelli has a more southern and fragmented distribution. In this study, we assessed whether nuclear markers reveal similar evolutionary patterns to those inferred from mtDNA variation. We studied a battery of allozyme and microsatellite loci in a geographically representative set of individuals from both species. For each species we evaluated overall levels of differentiation, patterns of geographical variation in genetic diversity, genetic relationships amongst localities, and applied model-based individual multilocus genotype clustering approaches to detect hidden population structure. Our results for P. bocagei are highly concordant with the phylogeographical scenario inferred from mtDNA variation: we found very low levels of population differentiation, consistent with survival in a single glacial refugium, and detected signatures of a rapid demographic and geographical expansion. The analyses of nuclear markers furthermore helped to identify a probable refugial area, as well as expansion routes. Additionally, in concordance with observations based on mtDNA variation, a low level of population differentiation was observed in P. carbonelli, but this was significantly higher than in P. bocagei. However, the geographical basis for differentiation in P. carbonelli is highly inconsistent between mtDNA and nuclear markers, suggesting a complex, albeit recent, history of fragmentation. A recent reduction of this species’ distribution has probably erased the signatures of glacial isolation and post-glacial expansion that are normally found in other Iberian species, suggesting that the currently observed pattern of genetic differentiation in this species was shaped more by recent genetic drift than by the Pleistocene climatic oscillations.

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We report new primers for the amplification and sequencing of 11 nuclear markers in squamate reptiles and anuran amphibians (five in squamates, six in anurans). Ten out of the 11 loci are introns (three of which are linked) that were amplified using an exon-primed, intron-crossing (EPIC) PCR strategy, whereas an eleventh locus spans part of a protein-coding gene. Squamate and anuran primers were initially developed for Lacerta schreiberi (Squamata: Lacertidae) and Pelodytes spp. (Anura: Pelodytidae), respectively. Cross-species amplification of the squamate markers was evaluated in four genera representing two additional families, whereas for anurans three genera corresponding to three additional families were tested. Three out of the five loci were successfully sequenced in all squamate taxa tested. Cross-amplification of the six anuran markers had lower, but still significant, success. We predict these markers will be of great utility for both population genetics and phylogenetic studies.

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

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The monophyletic species subgroup of Podarcis tauricus is distributed in the western and southern parts of the Balkans, and includes four species with unresolved and unstudied inter- and intra-specific phylogenetic relationships. Using sequence data from two mitochondrial and three nuclear genes and applying several phylogenetic methods and species delimitation approaches to an extensive dataset, we have reconstructed the phylogeny of the Podarcis wall lizards in the Balkans, and re-investigated the taxonomic status of the P. tauricus species subgroup. Multilocus analyses revealed that the aforementioned subgroup consists of five major clades, with P. melisellensis as its most basal taxon. Monophyly of P. tauricus sensu stricto is not supported, with one of the subspecies (P. t. ionicus) displaying great genetic diversity (hidden diversity or cryptic species). It comprises five, geographically distinct, subclades with genetic distances on the species level. Species delimitation approaches revealed nine species within the P. tauricus species subgroup (P. melisellensis, P. gaigeae, P. milensis, and six in the P. tauricus complex), underlining the necessity of taxonomic re-evaluation. We thus synonymize some previously recognized subspecies in this subgroup, elevate P. t. tauricus and P. g. gaigeae to the species level and suggest a distinct Albanian-Greek clade, provisionally named as the P. ionicus species complex. The latter clade comprises five unconfirmed candidate species that call for comprehensive studies in the future.

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Samplings carried out in the municipality of A Coruña, one of the smaller surface area of Galicia, showed the presence of 8 species of amphibians and 9 species of reptiles. Eight of them (4 amphibians and 4 reptiles) were relatively frequent and very sparse the rest. Some of these species can survive even in disturbed areas. However, alterations of the natural environment, such as eucalyptus plantations, abandonment of farmland and the presence of residues, can seriously affect the survival of these species.

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Giardia duodenalis is a zoonotic parasite that infects the gut of a wide range of vertebrates, including numerous wildlife species. However, little is known about this protozoan parasite in reptiles. Fecal samples from 31 wild lizards were collected in Galicia (northwest Spain) and screened for the presence of Giardia by PCR amplification and sequencing of the ITS1-5.8S-ITS2 region in the ribosomal unit. This allowed detection of the parasite in 5 samples (16.1%), and enabled identification of G. duodenalis assemblage A2 in two samples of Iberian rock lizard (Iberolacerta monticola), G. duodenalis assemblage B in other two samples of I. monticola, and G. duodenalis assemblage E in one sample of Bocage`s wall lizard (Podarcis bocagei). The results obtained after PCR amplification and sequencing of the SSU-rDNA gene confirmed the presence of G. duodenalis assemblage A in two samples of I. monticola. This is the first report of G. duodenalis in free-living lizards, although further studies are needed to distinguish between actual infection and mechanical dissemination of cysts. The detection of zoonotic and livestock-specific assemblages of G. duodenalis demonstrates the wide environmental contamination by this parasite, possibly due to human activities.

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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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Host parasite specificity is related to several factors including the type of life cycle of the parasite, the habitat occupied by both parasites and hosts, and the food habits of the hosts. In this work, gastrointestinal helminths in two congeneric lacertid lizards, Podarcis bocagei and Podarcis carbonelli living in sintopy, have been analysed. Out of four helminth species found, three of them, the trematode Brachylaima sp., the cestode Oochoristica gallica and the nematode Skrjabinelazia hoffmanni, have indirect life cycles while the nematode Spauligodon carbonelli has a direct life cycle. Heteroxenous helminths showed low host specificity whereas S. carbonelli may be placed at an intermediate position in a continuum of the degree of specificity.

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A parasitological survey was carried out to determine the relationships between helminth fauna and some biological traits of two host species of lacertid lizards. Size, sex, diet, and season of collection, as well as infracommunities and component communities of Podarcis bocagei and P. carbonelli from Douro litoral (northwest Portugal), have been analysed. In both host species, we found low values of parasite infection parameters and diversity. These findings were in accordance with their feeding habits, eating only animal prey and no plant matter. Evidences suggest a relationship between the presence of Brachylaima sp (Trematoda: Brachylaimidae) and the ingestion of snails by both lizard species. Spauligodon carbonelli (Nematoda, Pharyngodonidae), a specialist in Podarcis hosts, was the most important parasite, affecting mainly adult males with a prevalence of infection showing seasonal variation.

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The presence of Haemogregarines (sensu lato) in the blood of many species of reptiles is not at all unusual. In Iberian and Canarian lacertid lizards, Haemogregarines have been recorded infecting erythrocytes. Most of the records were the mature gametocytes, and this fact always has disabled the accurate identification of these infecting forms because their morphology do not give enough information to identify them even at generic level. We have analyzed blood smears from 75 specimens of Podacis bocagei (Seoane, 1884) and 33 specimens of Podarcis carbonelli (Pérez- Mellado, 1981) from several localities of North-East Portugal. We found Haemogregarines (sensu lato) for 74.7% of P. bocagei and 69.7% of P. carbonelli. In order to progress in the identification of these Haemogregarines, we studied some characteristics other than the morphology of the mature gametocytes. Thus, we observed immature gametocytes (considered merozoytes by some authors) oval in form, and with a big central nucleus. Furthermore, we analyzed histologically the liver of 4 of the searched hosts (2 P. bocagei, 2 P. carbonelli). We detected the presence of latent cysts with sporozoytes. Both characteristics, the presence of merozoytes in the blood and the presence of cysts in the liver, have been recorded by several authors as typical of the genus Hepatozoon. Although more histological analysis are necessary to assert that the hole of the sample is infected by Hepatozoon, we suspect that not only P. bocagei and P. carbonelli from Portugal, but many species of Iberian (and probably Balearic and Canary) lacertid lizards (on which we are now developing some investigations) could be infected by species of this genus.

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In Iberian and Canarian lizards, haemogregarines have been recorded infecting erythrocytes, but most of the records correspond to mature gametocytes. We analysed blood smears from 75 specimens of Podarcis bocagei (Seoane) and 33 specimens of P. carbonelli (Pérez-Mellado) from localities of north-western Portugal. We found haemogregarines in 74.7% of P. bocagei and 69.7% of P. carbonelli. Our observations show characteristics of the haemogregarines other than the morphology of the mature gametocytes. In histological sections of the liver of four hosts latent cysts with sporozoites and meronts with merozoites were detected. Both traits have been described as typical of the genera Hepatozoon Miller, 1908 and Hemolivia Petit, Landau, Baccam & Lainson, 1990. We suggest that not only P. bocagei and P. carbonelli from Portugal but other species of Iberian and Canarian lacertids might also be infected by species belonging to one or both genera.

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The geographic distributions of the Bocage’s wall lizard, Podarcis bocagei and the Carbonell’s wall lizard, P. carbonellaein Portugal were determined through extensive field surveys. The river Douro constitutes the main geographical barrier where both lizard species show nearby a zone of contact. Predictive models of probability of occurrence were developed for both species in Portugal, based on the Logistic Multiple Regres- sion (LMR) and the Overlap Analysis (OA). On a coarse scale, the best-fit model suggested that the distribution of P. bocagei in Portugal might be largely explained by macroenvironmental variables such as radiation, eva- potranspiration or type of climate. The balance between the number of frost days per year and a coefficient of aridity (Kdr = 100 × temperature/precipitation) seems important to explain the range of P. carbonellae. Howe- ver, the large overlap found between predictive areas strongly suggest that historical events (e.g. Pliocene spe- ciation, Pleistocene glacial refugia) might have been preponderant, acting the macroenvironmental factors as ultimate determinants of the present-day distribution of both species. The distribution of P. bocagei is signi- ficantly congruent with that of Chioglossa lusitanica, partially with Vipera seoaneiand with Rana ibericaand Lacerta schreiberinorthwards the river Douro. Conversely, the contractive distribution ofP. carbonellaedid not share any significant pattern of congruence with the latter taxa.

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Six biometric characters and 15 electrophoretic loci of three known subspecies of Podarcis bocagei were studied. Contrary to biometric indications, Podarcis bocagei berlengensis showed a closer genetic relationship with Podarcis bocagei carbonelli than with the nominal subspecies. The biometric results were confounded by the relative large size of P. b. berlengensis. Post-glacial colonization from two different areas might explain the greater genetic differentiation between the P. b. bocagei and P. b. carbonelli populations (Nei`s D - 0.158 - near the species threshold). It might also explain the known colonization of these two lizards through the islands off the west coast of the Iberian penninsula and their present distributions to the north of (Galicia; Northern Portugal) and to the south of the valley of river Douro (central Portugal, western and Sistema Central), respectively.

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The present study reports the morphology of leukocytes of 12 European lacertid lizards (Podarcis sicula, P. tiliguerta, P. melisellensis, P. bocagei, P. muralis, Algyroides nigropunctatus, Lacerta viridis, L. bilineata, L. trilineata, L. oxycephala, Timon lepidus, and Zootoca vivipara) stained using May–Grünwald/Giemsa method. The morphology of white blood cells was very similar among species, suggesting a relative morphological uniformity within the lacertid lizards. For six species (i.e. P. sicula, P. tiliguerta, P. melisellensis, P. bocagei, P. muralis, and A. nigropunctatus), we determined the leukocyte differential counts, which may be considered representative of the normal values of the corresponding populations. These results may be useful either in clinical investigation to detect pathologies in wild individuals, as in management and conservation projects to assess the general health conditions of natural wild lizard populations.

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Podarcis bocagei and P. guadarramae are sister taxa overlapping geographically in north-western Spain and northern Portugal. These two species are often found in syntopy, although they differ in morphology and habitat use. In particular, P. guadarramae is characterized by a flat body, possibly associated with more saxicolous habits with respect to P. bocagei, a ground-dwelling species. From a physical point of view, it can be hypothesized that P. guadarramae will lose more water than P. bocagei, given the higher surface to volume ratio of its body. Alternatively, since P. bocagei is restricted to Atlantic climate while P. guadarramae may also occupy Mediterranean areas, it can be hypothesized that the second might be more resistant to water loss and prefer higher body temperature. During summer, we collected adult male lizards in Moledo, northern Portugal, where the two species are syntopic. Lizards were tested in thermal gradients and water loss was assessed using incubators with three temperature treatments (27, 32 and 37 °C). Results suggested that the two species only slightly differ in thermal preferences. However, when exposed to different thermal regimes, they lost water at different rate. In particular, while at 27 °C the two species are similar, P. guadarramae is more resistant to water loss at higher temperatures. Our experiments did not support a purely physical hypothesis. The results corroborate the hypothesis that in these two species there has been physiological divergence in resistance to water loss. In comparison with P. bocagei, P. guadarramae might live in more arid habitats and be associated with bigger rocks and more exposed surfaces when in syntopy. Ecological differences between these two species are at least partly associated with physiological resistance in water loss. In contrast, the absence of a neat difference in thermal preferences might suggest that preferred temperature is a conservative trait in these sister taxa.

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Sister species living in sympatry offer the opportunity to study the degree of divergence in their ecological, physiological and life-history traits. It has been hypothesized that closely related species with overlapping distribution should differ in their niche to reduce competition for resources. Furthermore, the investigation of sympatric species may shed light on how they may coexist without outcompeting each other. In the present study, we assess the degree of physiological divergence in two sympatric lacertid lizards, Podarcis bocagei and Podarcis guadarramae lusitanicus. These species share a Pliocenic ancestry and overlap at a both geographical and ecological scale. We assessed their thermal preferences and water loss rates, two physiological traits considered stable across congeneric species. We found that the two species differ in both traits, with P. bocagei selecting higher temperature than P. g. lusitanicus and losing more water than the latter at and above its preferred temperature. The results also showed that for both species body size has a relevant impact on thermal and hydric traits, with bigger individuals losing proportionally less water and selecting higher temperatures. These results, combined with previous evidence, suggest that physiological mechanisms, ecological preferences and morphology probably allow these two species to overlap in their distribution while selecting different microhabitats and thus decreasing possible competition between them.

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Climate change is negatively affecting many species. The increase in mean air temperature is often associated with shifts in distribution, changes in phenology, and local extinctions. Other factors that only partially correlate with air temperature, like water shortage, may also contribute to the negative consequences of climate change. Although the effect of temperature on lizards’ ecophysiology is highly studied, many lizards are also at risks of increased water loss and dehydration, which are predicted to increase under climate change. Here we aimed for the first time to explore if lacertid lizards exposed to dehydration thermoregulate less precisely than hydrated lizards and if dehydrated lizards are less active, change the daily pattern of thermoregulation and balance water balance against thermoregulation. We exposed four lizard species with differences in the thermal preference to thermal gradients with or without a source of water. We measured preferred body temperatures, daily pattern of thermoregulation, and the use of space. Dehydration negatively affected thermoregulation in all investigated species. Dehydrated lizards reduced their preferred body temperature and showed a species-specific pattern of hourly change in thermal preference. Furthermore, they more frequently used the colder parts of the gradients and spent more time hidden. Lizards experiencing dehydration may suffer a reduction in survival and fitness because of poor thermoregulation. Similarly, they may spend more time hidden, waiting for more favourable weather conditions. Such inactivity may carry ecological costs especially in those regions that undergo either short or prolonged periods of droughts.

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Although the spatial context is expected to be a major influence in the interactions among organisms and their environment, it is commonly ignored in ecological studies. This study is part of an investigation on home ranges and their influence in the escape behaviour of Iberian lizards. Fieldwork was conducted inside a 400 m2 mesocosm, using three acclimatized adult male individuals. In order to perform analyses at this local scale, tools with high spatial accuracy are needed. A total of 3016 GPS points were recorded and processed into a Digital Elevation Model (DEM), with a pixel resolution of 2 cm. Then, 1156 aerial photos were taken and processed to create an orthophoto. A refuge map, containing possible locations for retreats was generated with supervised image classification algorithms, obtaining four classes (refuges, vegetation, bare soil and organic soil). Furthermore, 50 data-loggers were randomly placed, recording evenly through the area temperature and humidity every 15’. After a month of recording, all environmental variables were interpolated using Kriging. The study area presented an irregular elevation. The humidity varied according to the topography and the temperature presented a West-East pattern. Both variables are of paramount importance for lizard activity and performance. In a predation risk scenario, a lizard located in a temperature close to its thermal optimum will be able to escape more efficiently. Integration of such ecologically relevant elements in a spatial context exemplifies how remote sensing tools can contribute to improve inference in behavioural ecology.

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Lizards use flight as main anti-predator behaviour, frequently escaping to a refuge. This response is well studied in terms of morphology, physiology and reproductive biology, though the spatial context is frequently ignored. The home range influence or the spatial factors of refuge selection are not completely understood. This study aims to determine the influence of Podarcis bocagei home range on its escape behaviour and how is it established when facing a new area. This study was conducted inside a 400m2 mesocosmos using 38 acclimatized adult individuals. Remote sense techniques were used in order to map the mesocosm, which resulted in an orthophoto, temperature and humidity map and refuge map. The individuals were approached with a constant pace and manually marked three georeferenced points in the orthophoto (predator location, starting and final escape locations). We estimated home-ranges using a 90% Characteristic Hull Polygon. This study revealed the home range is gradually established. At first, the individuals roam randomly through the mesocosm in an exploratory phase and then established a home range that decreased with the time. About the escape outside the home range, 25% of the escapes ended outside the home-range, although keeping short distance (about 50 cm). Lizards may temporally flee outside home range limits, but keeping a short distance allowing them to easily return, and hence, ensuring the mid-term persistence of their home ranges.

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The Pyrenees, Sierra Nevada, Sierra de Peña de Francia and the Cantabrian Mountains were visited during the second half of May, 1994. In northern Spain Podarcis hispanica and P. muralis are living in syntopy. On all sites Lacerta lepida was common. North of Murcia the only typical L. lepida nevadensis have been seen. In the Sierra Nevada this subspecies had a large amount of green colouration. South of Madrid Acanthodactylus erythrurus and Psammodromus algirus were found in nearly every habitat. In the vicinity of Béjar Podarcis bocagei and Lacerta schreiberi were seen. In the Sierra Covadonga a large population of L. schreiberi was found in a very dry mountain slope.

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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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Species can occupy different realised niches when sharing the space with other congeneric species or when living in allopatry. Ecological niche models are powerful tools to analyse species niches and their changes over time and space. Analysing how species’ realised niches shift is paramount in ecology. Here, we examine the ecological realised niche of three species of wall lizards in six study areas: three areas where each species occurs alone; and three areas where they occur together in pairs. We compared the species’ realised niches and how they vary depending on species’ coexistence, by quantifying niche overlap between pairs of species or populations with the R package ecospat. For this, we considered three environmental variables (temperature, humidity, and wind speed) recorded at each lizard re-sighting location. Realised niches were very similar when comparing syntopic species occurring in the same study area. However, realised niches differed when comparing conspecific populations across areas. In each of the three areas of syntopy, the less abundant species shift its realised niche. Our study demonstrates that sympatry may shift species’ realised niche.

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The analysis of the spatial structure of animal communities requires spatial data to determine the distribution of individuals and their limiting factors. New technologies like very precise GPS as well as satellite imagery and aerial photographs of very high spatial resolution are now available. Data from airborne LiDAR (Light Detection and Ranging) sensors can provide digital models of ground and vegetation surfaces with pixel sizes of less than 1 m. We present the first study in terrestrial herpetology using LiDAR data. We aim to identify the spatial patterns of a community of four species of lizards (Lacerta schreiberi, Timon lepidus, Podarcis bocagei, and P. hispanica), and to determine how the habitat is influencing the distribution of the species spatially. The study area is located in Northern Portugal. The position of each lizard was recorded during 16 surveys of 1 h with a very precise GPS (error < 1 m). LiDAR data provided digital models of surface, terrain, and normalised height. From these data, we derived slope, ruggedness, orientation, and hill-shading variables. We applied spatial statistics to determine the spatial structure of the community. We computed Maxent ecological niche models to determine the importance of environmental variables. The community and its species presented a clustered distribution. We identified 14 clusters, composed of 1–3 species. Species records showed two distribution patterns, with clusters associated with steep and flat areas. Cluster outliers had the same patterns. Juveniles and subadults were associated with areas of low quality, while sexes used space in similar ways. Maxent models identified suitable habitats across the study area for two species and in the flat areas for the other two species. LiDAR allowed us to understand the local distributions of a lizard community. Remotely sensed data and LiDAR are giving new insights into the study of species ecology. Images of higher spatial resolutions are necessary to map important factors such as refuges.

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Home ranges in animals can be estimated by different methods like minimum convexpolygons, characteristic hulls or kernels while correlative ecological niche models(ENMs) are commonly employed for forecasting species’ranges. However, ENMscan also model the distribution of individuals if environmental very high spatial reso-lution data are available. Indeed, remote sensing (RS) can provide images with pixelsizes of few centimetres. Here, we modelled the distribution of individual lizards(Podarcis bocagei) combining aerial-like photographs recorded with a compact cam-era and a matrix of temperature/humidity data-loggers to obtain several environmen-tal layers with very high spatial resolution. We recorded lizards’positions in a20920 m mesocosm with a high precision GPS device (~10 cm of error), multipletimes per day throughout the whole period of daily activity. We built an orthophotomap (pixels of 20 cm2) from camera pictures, a digital surface model, and a land-cover supervised classification map. We recreated climate-like variables by combin-ing data-logger data. For each individual, we calculated the distance to males andfemales, excluding the focal lizard. We computed individual realized niche modelswith Bioclim, GAM, GLM, Maxent and random forest. Models attained a very highevaluation score in most cases. The most contributing variables were related tomicroclimate (isothermality, minimum temperature and humidity) and distance toconspecifics. Our very high spatial resolution models provided robust information onhow space is used by each lizard. Correlative models can identify the most suitableareas inside the home range, similar to core areas estimated from kernel algorithms,but allowed better statistical inference. Overall, RS tools generated high-quality envi-ronmental data, and when combined with ENMs, improved the robustness of the pre-dictions on spatial patterns of small terrestrial animals.

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There is an increasing amount of evidence pointing towards the importance of bacteria and parasites in the ecology and dynamics of host populations. Complex microorganism communities living in intestinal tracts of the animals are known as the gut microbiome. These bacterial communities can impact the host’s fitness, immune system and behavior, whereas microbiome composition and diversity can be influenced by countless factors such as the host’s taxonomy, habitat, diet and social interactions with other individuals. In turn, blood parasites have a fundamental role in the ecosystem, weakening host fitness and reproductive ability, and thereby influencing population dynamics. Five lacertid lizard species were used for the present work, three endemic and two introduced: in Parque das Nações, Lisbon, the invasive Podarcis siculus (n=20) and the native Podarcis virescens (n=22); another introduced species, Teira dugesii (n=20), was captured from a distinct area of Lisbon (9 kilometres from the Parque das Naçoes); and in Moledo, Northern Portugal, two endemic and syntopic species were studied – Podarcis bocagei (n=33) and Podarcis lusitanicus (n=8). This thesis comprises two investigations. A first study was conducted in order to better understand the role and influence of environmental factors and the host in the gut microbiome composition and diversity of these five species. The second study had the objective of evaluating the effect of blood parasites on host behavior (using a mirror for the behavioral test), with the two syntopic species found in Moledo as models. This project allowed me to characterize the gut microbiome of these five species for the first time. Habitat/locality were the main factors driving gut microbiome differences between these species, and there was a greater microbiome diversity in species captured in Lisbon, the most urbanized studied site. I was able to verify a positive correlation between bacterial diversity and size in P. siculus, which indicates that it accumulates more bacteria, likely because bigger lizards explore more diverse environments. Additionally, I estimated bacteria transmission between sympatric lizards at Moledo and Parque das Nações. High transmission was verified for both populations. Results from the Lisbon population indicate that the invasive species, P. siculus might have acquired a higher proportion of local microbiota through the native species, P. virescens than vice versa. The second study highlighted the infection prevalence differences between P. bocagei and P. lusitanicus, with a larger number of infected individuals in the former. However, P. lusitanicus registered a higher average parasite intensity. The species that interacted with the mirror the most, and was therefore considered bolder, was P. lusitanicus. Contrary to what was expected, the individuals that interacted with the mirror the most, were more parasitized. This thesis aids in understanding the behavior and broadening the sphere of knowledge of microorganisms (bacteria and microparasites) and their interactions with the host. Such work is fundamentally important, and increased investment in these areas of knowledge is essential for species conservation.

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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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Host-gut microbiota interactions are complex and can have a profound impact on the ecology and evolution of both counterparts. Several host traits such as taxonomy, diet and social behavior, and external factors such as prey availability and local environment are known to influence the composition and diversity of the gut microbiota. In this study, we investigated the influence of taxonomy, sex, host size, locality/habitat on gut microbiota diversity in five lizard species from two different sites in Portugal. We also analyzed the potential levels of microbial transmission between species that live in sympatry and syntopy. We studied Podarcis bocagei and Podarcis lusitanicus from northern Portugal (Moledo); and two invasive species, Podarcis siculus and Teira dugesii, and the native Podarcis virescens from Lisbon. We used a metabarcoding approach to characterize the bacterial communities from the cloaca of lizards, sequencing the V4 region of the 16S rRNA. Habitat/locality was found to be the main driver of the differences in composition and structure of gut bacterial communities of the studied lizards, with host effects more evident at finer taxonomic scales. Additionally, lizards from urbanized environments had higher microbiome diversity than lizards from rural areas. We detected a significant positive correlation between size and gut bacterial alpha-diversity in the invasive species P. siculus, which could be due to higher exploratory behaviours. Moreover, estimates of bacterial transmission indicate that P. siculus may have acquired a high proportion of local microbiota. These findings indicate that a diverse array of host and environmental factors can influence lizards gut microbiota.

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Host-gut microbiota interactions are complex and can have a profound impact on the ecology and evolution of both counterparts. Several host traits such as systematics, diet and social behavior, and external factors such as prey availability and local environment are known to influence the composition and diversity of the gut microbiota. In this study, we investigate the influence of systematics, sex, host size, and locality/habitat on gut microbiota diversity in five lizard species from two different sites in Portugal: Podarcis bocagei and Podarcis lusitanicus, living in syntopy in a rural area in northern Portugal (Moledo); the invasive Podarcis siculus and the native Podarcis virescens, living in sympatry in urbanized environment (Lisbon); and the invasive Teira dugesii also living in an urban area (Lisbon). We also infer the potential microbial transmission occurring between species living in sympatry and syntopy. To accomplish these goals, we use a metabarcoding approach to characterize the bacterial communities from the cloaca of lizards, sequencing the V4 region of the 16S rRNA. Habitat/locality was an important factor explaining differences in gut bacterial composition and structure, with species from urbanized environments having higher bacterial diversity. Host systematics (i.e. species) influenced gut bacterial community structure only in lizards from the urbanized environment. We also detected a significant positive correlation between lizard size and gut bacterial alpha-diversity in the invasive species P. siculus, which could be due to its higher exploratory behavior. Moreover, estimates of bacterial transmission indicate that P. siculus may have acquired a high proportion of local microbiota after its introduction. These findings confirm that a diverse array of host and environmental factors can influence lizards gut microbiota.

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This paper is divided into two parts: classic knowledge on theory of island biogeography (I) and the evolution of behaviour and morphology under insular constraints (II). The main subjects dealt with are: the problem of small populations (1); the biological definition of an island and the importance of biological studies on islands (2); the species-area effect and its consequences for a nature conservation strategy (3); the equilibrium theory and the weaknesses of this model (4); the founder principle (5); some features of insular populations, such as clutch size, niche enlargement and changes in body size (6); the natural history of colonisation and the success and failure of colonisation (7); the morphological paradigm of island faunas (8) and the evolution of social behaviour under insular constraints – territoriality, altruism, social structure and its relationships with body shape (9).