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Algyroides fitzingeri, Algyroides marchi, Algyroides moreoticus, Algyroides nigropunctatus, Lacerta agilis, Lacerta lepida, Lacerta princeps, Lacerta schreiberi, Lacerta strigata, Lacerta trilineata, Lacerta viridis, Lacerta andreanszkyi, Lacerta armeniaca, Lacerta bedriagae, Lacerta brandtii, Lacerta cappadocica, Lacerta caucasica, Lacerta chlorogaster, Lacerta cyanura, Lacerta danfordi, Lacerta derjugini, Lacerta dugesii, Lacerta fraasii, Lacerta graeca, Lacerta horvathi, Lacerta jayakari, Lacerta laevis, Lacerta monticola, Lacerta mosorensis, Lacerta oxycephala, Lacerta parva, Lacerta perspicillata, Lacerta praticola, Lacerta rudis, Lacerta saxicola, Lacerta vivipara, Gallotia atlantica, Gallotia galloti, Gallotia simonyi, Podarcis erhardii, Podarcis filfolensis, Podarcis hispanica, Podarcis lilfordi, Podarcis melisellensis, Podarcis milensis, Podarcis muralis, Podarcis peloponnesiaca, Podarcis pityusensis, Podarcis sicula, Podarcis taurica, Podarcis tiliguerta, Podarcis wagleriana, Psammodromus algirus, Psammodromus blanci, Psammodromus hispanicus, Psammodromus microdactylus.

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The application of molecular markers to the study of geographic variation in widely distributed species has often revealed new species that had been overlooked because of their superficial resemblance to related taxa. A good example are lizards in the Podarcis hispanica s.l. complex, which after taxonomic revision (Geniez et al., 2014) was split into several species. In this communication we focus on two of them, Podarcis guadarramae and P. virescens, broadly distributed in central Iberia but with poorly defined ranges in Comunidad de Madrid and neighboring regions. García París et al. (1989) first described the distribution of Podarcis hispanica s.l. in Madrid, illustrating with pictures distinctive coloration patterns associated to different areas in the region, but since then there have been no followup studies taking into account taxonomic changes and delineating the ranges of the two species in the complex that are present in Madrid. In this study we delimited the range boundaries of P. guadarramae and P. virescens in Comunidad de Madrid through a combination of molecular analyses and morphological data and the modeling of their potential distributions. The occurrence of Podarcis guadarramae in Madrid comprises the mountains of Sierra de Guadarrama and their foothills, on geological settings of quartzite, granite, and shale, at altitudes ranging from 650 m in the Alberche river valley up to 2000 m in the southern slopes of Siete Picos, with an incompletely defined contact zone with Podarcis virescens extending parallel to Sierra de Guadarrama, along the municipalities of Cenicientos, San Martín de Valdeiglesias, Villamanta, Zarzalejo, Fresnedillas de la Oliva, Torrelodones, Hoyo de Manzanares, Soto del Real, El Berrueco and El Cardoso. We confirm the record of P. guadarramae in Torrejón de la Calzada, and identify other population isolates in the city of Toledo (confirmed with molecular data) and San Martín de Montalbán (morphological data). Podarcis virescens spreads through most of the Madrid region, from 494 m in Aranjuez up to around 950 m in Colmenar Viejo, not overlapping with P. guadarramae, thus displaying a complementary distribution over geological settings of gravel, sandstone, and dolomites in the southeastern part of the region. This species is common in constructions and anthropized areas, and can be found mostly in low elevation plains and plateaus, often in agricultural land, but also in urban areas and parks, as well as in natural habitats like forested hills with rocky outcrops. Our results suggest the possible existence of ecological differentiation processes that might contribute to reproductive isolation between the two species.

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Under a documented paleogeographic history of the Ria de Arosa archipelago, northwestern Spain, predictions can be made on the duration and strength of isolation of Podarcis guadarramae lizard populations that were ‘captured’ on the islands when sea-levels rose. We predict that genetic diversity: i) is lower on islands than on the mainland, ii) increases with island size, and iii) decreases with increasing distance from the mainland. We also predict that the population genetic structure across islands and the mainland conforms to a pattern of isolation by distance (iv). To test these hypotheses, we analysed a panel of nine highly polymorphic microsatellite loci. The first (i) hypothesis was corroborated. We discuss our findings in the light of the effects of habitat fragmentation and lizard conservation.

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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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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 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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Reptiles and amphibians in Portugal The author spend two weeks in July 2002 in Beira, Portugal. The following observations were made: Vale de Asna (Serra do Açor): the sound of several Alytes obstreticans boscai (probably), several Tarentola mauritanica, one Podarcis hispanica type 1, many Psammodromus algirus, several Chalcides chalcides striatus, larvae of Rana iberica, Triturus marmoratus marmoratus, Triturus helveticus sequeirai and Triturus boscai, one adult Rana perezi, one dead Bufo bufo, three Natrix maura (half-grown and adult). Waterfall of Sobral (Serra do Açor): many Psam-modromus algirus, several larvae, juvenile and adult Rana iberica, many juvenile Bufo bufo. Pena (Serra da Lousã): several Psammodromus algirus. Mata da Margaraça (Serra do Açor): several Psam-modromus algirus, one Podarcis hispanica (probably). Fraga de Pena (Serra do Açor): several larvae and two adult Rana iberica, one Natrix maura (probably), several larvae and three adult Chioglossa lusitanica. Serra da Estrela: Near the top of mount Torre (between 1750 and 1993 m): many Iberolacerta monticola monticola, many larvae of Bufo species, many adult Rana perezi, one adult Rana iberica, one adult Hyla arborea molleri. Around 1200 m, one adult and one juvenile Timon lepidus, one adult Lacerta schreiberi. At 1600 m one juvenile Lacerta schreiberi. Between Piódão and Foz d’Égua (Serra do Açor): one half-grown Natrix natrix astreptophora, one juvenile Natrix maura (probably newly hatched), several Psammodromus algirus and one adult Malpolon monspessulanus. It is stated that Portugal is a very interesting place for spotting reptiles and amphibians in nature.

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

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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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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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Investigadores del Museo Nacional de Ciencias Naturales (MNCN-CSIC) han analizado la respuesta a las condiciones ambientales de la lagartija de Guadarrama, Podarcis guadarramae, perteneciente al complejo de especies de la lagartija ibérica, y han descubierto que las diferencias morfológicas entre poblaciones están determinadas por las condiciones ambientales que experimentan más que por sus diferencias genéticas.

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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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One of Darwin`s most widely known conjectures is that prey are tame on remote islands, where mammalian predators are absent. Many species appear to permit close approach on such islands, but no comparative studies have demonstrated reduced wariness quantified as flight initiation distance (FID; i.e. predator–prey distance when the prey begins to flee) in comparison with mainland relatives. We used the phylogenetic comparative method to assess influence of distance from the mainland and island area on FID of 66 lizard species. Because body size and predator approach speed affect predation risk, we included these as independent variables. Multiple regression showed that FID decreases as distance from mainland increases and is shorter in island than mainland populations. Although FID increased as area increased in some models, collinearity made it difficult to separate effects of area from distance and island occupancy. FID increases as SVL increases and approach speed increases; these effects are statistically independent of effects of distance to mainland and island occupancy. Ordinary least-squares models fit the data better than phylogenetic regressions, indicating little or no phylogenetic signal in residual FID after accounting for the independent variables. Our results demonstrate that island tameness is a real phenomenon in lizards.

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A population of the lacertid lizard Podarcis hispanica was studied in the Sistema Central Mountains in Salamanca Province, Spain. Lizards confined their activities to stable home ranges, but size range varied enormously between individuals. Home ranges generally contained 1) a high proportion of rocks > 100 cm in height and 2) a significantly higher abundance of crevices than expected. The overall abundance of lizards increased with decreasing amounts of shrubs and herbaceous plants cover and densities were highest in bare and rocky areas. Adult females selected particular habitat types according to the season, whereas habitat selection of males and juveniles did not differ between spring and summer. Non-resident adult lizards and juveniles occupied habitats of marginal quality in relation to crevice abundance.

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In this work we studied the effect of an experimental increment of density on survival of the Iberian wall lizard, Podarcis hispanica. The manipulation of adult males at the experimental plat located in the western zone of the Sistema Central (Iberian Peninsula). In our study area, P. hispanica shows one of the highest densities reported for Podarcis genus in the mainland. Densities not experimentally altered varied between 530 ha-1 and 920 ha-1, excluding hatchlings (Jolly-Seber model estimates). A departure from 1:1 sex-ratio was observed in the population under study, adult females being more numerous than adult males. Survival probability of adult males, residents and introduced, greatly decreased after the introduction of adult males at the experimental plot, but survivorship of adult females and subadult individuals of both sexes was not affected. Reduction in survival probability of resident and introduced males was directly related to density of introduced males. However, we did not detect any relation between their survival probability and density of resident adult males. Density could influence survival through competition. Thus, increment of density had a significant effect on frequency of aggressive encounters among adult males, but not on other age and sex categories. In spite of the fact that subadult individuals were not affected by the introduction of adult males at the experimental plot, their survival probability was significantly reduced when density of resident adult males increased, perhaps because of an asymmetric competition or interference from older individuals. In conclusion, our experimental increment of adult males density has evidenced a balanced demographic structure in relation to environmental resources, and that density does not have the same effect with respect to age and sex.

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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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Monfragüe National Park (Cáceres, Extremadura, Spain) is a protected area in central‐western Iberia, including some of the best preserved primary Mediterranean vegetation. Legal protection dates back to 1979 (first as a Natural Park and then as a National Park), but knowledge about its reptile communities is so far limited to sparse records. In this paper we present an updated species list based on 521 records covering 163 1x1 km UTM grids in the study area, compiled in the period 2000‐2019. We detected 20 native species, representing 71.4% of the reptile fauna in Extremadura and 35% of the ibero‐balearic reptile fauna. Additionally, based on molecular analyses populations of the amphisbaenid genus Blanus in the study area are assigned to the oriental Iberian taxon, B. cinereus. The new records extend the known distribution of the different reptile species in the study area in 56 10x10 km UTM grids. Species presenting more restricted distributions in Monfragüe are Lacerta schreiberi (one 1x1 grid), Emys orbicularis, and Acanthodactylus erythrurus (four 10x10 grids each).

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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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Socioeconomic and climatic factors are modifying fire regimes with an increase of fire frequency and extension. Unfortunately, the effects of recurrent fires on biological processes that ultimately affect the genetic diversity of animal populations are mostly unknown. We examined genetic patterns of diversity in the wall lizard Podarcis guadarramae in northern Portugal, one of the European regions with the highest percentage of burnt land. This species is a small saxicolous lizard as it inhabits natural outcrops and artificial stone walls, likely in recurrent-fire landscapes. We genotyped nine microsatellites from ten populations selected according to a gradient in fire recurrence, and compared genetic diversity indexes and demographic patterns among them. At the population level, we hypothesize that a high level of mortality and population bottlenecks are expected to reduce genetic heterozygosity in sampled localities affected by recurrent fires. Alternatively, genetic signatures are expected to be absent whether fire did not cause high mortality. Regardless of levels of mortality, we expect a gain in genetic diversity whether recurrent fires facilitate lizard dispersal and migration due to the increased quality of the habitat for wall lizards. At the regional level, we examine whether a recurrent fire regime may disrupt the spatial structure of populations. Our results showed an increase in genetic diversity in recurrently burnt populations, and a decline in longer-unburnt populations. We did not detect bottleneck effects in repeatedly-burnt populations. High genetic diversity in recurrent fire populations suggests a high dispersion rate between adjacent metapopulations and perhaps immigration from outside the fire boundary. At the regional level, lizard populations show low differentiation and weak genetic structure, suggesting no effects of fire. This study confirms field-based censuses showing that recurrent-fire regimes give ecological opportunities to wall lizards that benefit from habitat openness.

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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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There is an increasing recognition that fire is an important component of many ecosystems in the world, essential to understand the ecology and diversity of communities in fire-prone regions. It impacts fauna directly, through mortality, or indirectly by changing habitat structure and resources availability. In recent decades global warming, land abandonment and changes in traditional agricultural activities are driving changes in the frequency and extent of fires. In this context, to know the response of organisms to fire regimes is a conservation priority. Reptiles are a good model group to examine their responses to fire as they are ectothermic, sensible to modifications in habitat attributes such as vegetation structure and canopy closure, have low mobility and dispersal rates, and experience high rates of population declines and extinction. The purpose of this study is to evaluate the effects of fire on reptile assemblages at two geographic scales: at a worldwide scale, where I examined patterns of reptile responses to fire considering the time since fire effect with a meta-analysis methodology (Chapter 1); at a local scale, where I evaluated the effect of wildfires on the reptile communities of the Serra da Estrela Natural Park, namely on the population densities, species richness (Chapters 2) and examined how the trophic ecology and interspecific interactions varied with fire using the Psammodrome lizard as a model species (Chapter 3). For this, I used a set of proper methodologies in order to collect and analyze all the data needed to perform this study (search scientific data for a meta-analysis; transects for reptile surveys followed by vegetation and habitat structure characterization; fecal pellets analysis and network metrics for lizard‟s diet). At a worldwide scale, lizards‟ diversity (richness) but not abundance decreases immediately after fire (<1 year) but recovers quickly to pre-fire levels with increased abundance. Regarding snakes, diversity is not significantly affected by time-since-fire but abundance decreases in the latter stages of the post-fire succession. In Serra da Estrela Natural Park, reptile abundance decreases in the first 5 years of post-fire succession, but increases in the interval between 5-10 years. I also found that Psammodromus algirus is positively related with tree cover, and negatively related with herbaceous cover. Plant diversity is also a relevant parameter for reptiles, since its presence (abundance) is positively related with the diversity of plant species. Relatively to the diet, metrics of the bipartite network for lizards collected in unburned and burned habitats demonstrated that P. algirus presents more selective behaviors in unburned habitats, than in burned ones. I have also found differences between burned and unburned areas as regards availability of prey, being the burned area the one presenting the richest arthropod communities. My results suggest that immediately after fire there is a decrease in species richness but communities recover quickly and reptile abundance increases in early stages of post-fire succession (both globally, and also in Serra da Estrela Natural Park). Although reptiles present a close relation with habitat structure, they seem well adapted to areas with different time since fire and present stable abundances across the post-fire succession. Moreover, they have diet variation and are adapted to consume a large variety of preys when conditions are not suitable. In undisturbed areas, they may present a selective behavior.

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Interpopulational variation in sexual signals may lead to premating reproductive isolation and speciation. Genetic and morphological studies suggest that the Iberian wall lizard, Podarcis hispanica, forms part of a “species complex” with several cryptic species.We explored the role of chemical sexual signals in interpopulational recognition between five distinct populations of Iberian wall lizards in Central Spain. Results showed that these populations differed in morphology and in composition and proportion of chemical compounds in femoral gland secretions of males. Tongue-flick experiments indicated that male and female lizards discriminated and were more interested in scents of lizards from their own area (i.e., Northern versus Southern populations), but did not discriminate between all populations.Moreover, only males fromthe populations that are geographically located more far away preferred scent of females from their own population. These data suggest that, at least between some populations, there may be reproductive isolation mediated by chemical signals and cryptic speciation.

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In lizards, femoral gland secretions of males can have an important role in intraspecific communication. However, the chemosensory responses of lizards to specific compounds found in femoral secretions have been little studied. Steroids are the most abundant compounds in secretions, which have been shown to have a signalling function. However, other type of compounds might also be important. In this paper, we explored the role of alcohols in communication of Iberian wall lizards, Podarcis hispanica. We studied chemosensory behaviour, by measuring tongue flick rates, to test whether lizards could recognize several alcohols that are naturally found in femoral secretions. Results showed that lizards can discriminate and show chemosensory responses of different magnitude to different alcohols. The existence of high responses to alcohols suggest that, in addition to steroids, alcohols may also have a signalling function in femoral secretions of male P. hispanica lizards.

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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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Here I report on the reproductive characteristics of an insular population of the lacertid lizard Podarcis hispanica, which inhabits the Cíes Islands, an archipelago formed by three small islands off the northwest coast of Spain. The clutch characteristics of the population were investigated on the basis of clutches laid by pregnant females captured on these islands in 1999 and 2000. Other data were obtained principally by mark-recapture experiments. The egg-laying cycle of live females was determined by ventral palpation. Enlarged follicles were detected from March to July, and the laying period extended from April to the second half of July. The larger females initiated reproductive activities early in the season and produced up to three clutches annually, whereas the smaller individuals laid only a single clutch. Reproductive performance was related mainly to female size. Clutch size, individual egg mass, clutch mass, and hatchling size increased with the length of the females. I also examined the extent to which the reproductive characteristics of this insular population of P. hispanica diverged from those in other populations of the same species. I found no pronounced differences between these populations and other mainland populations of the same species of like body size. Several hypotheses may be proposed to explain the lack of major differences in reproductive characteristics (egg size and number). The first has to do with the relatively short length of time that this population has been isolated, and, therefore, it would still be too early for any divergence to have manifested itself. A second hypothesis is that the selection gradient for reproductive traits on the Cíes Islands may have remained similar to that on the mainland. A third hypothesis deals with the limitations imposed by the reduced body volume of this small-sized species on egg growth in conjunction with the shorter laying period of this population in an Atlantic climate.

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The traditional explanation of brain lateralization is that it avoids costly duplication of neural circuitry with the same function as well as decreasing the interference between different functions. This advantage could have important ecological implications in terms of predator–prey interactions. If species are lateralized in escape behavior at an individual level, any predator is facing individually asymmetric prey. On the other hand, if species are lateralized at a higher hierarchical level, such as at the population or species level, the predator can learn from experience arising from earlier encounters. In the present study, we investigate patterns of lateralization in 5 lizard populations of the Podarcis hispanica species complex, with the aim of analyzing the occurrence of lateralization at different hierarchical levels. Our results show that refuge selection during escape behavior is not universal in the P. hispanica species complex at different hierarchical levels, some individuals and populations are lateralized, with a consistent bias toward the refuge on the right. However, 2 populations showed no refuge preference, with lack of lateralization mainly arising due to the dominance of individuals with no preference rather than from a mix of right- and left-biased individuals. When considered at the P. hispanica species complex level, a pattern for right-refuge preference was found.

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

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

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Recent genetic works have suggested that the Iberian wall lizard Podarcis hispanicus (STEINDACHNER, 1870) sensu lato is a species complex. Several forms have already been elevated to species rank and linked to available nomina, but at least three still have to be formally named, including the western Iberian forms currently designated as Podarcis hispanicus“type 1A”, “type 1B” and “type 2”. The aim of the present work is to assign a valid nomen to these taxa. Using multivariate analyses, we first checked that the morphological differences reported in Portugal between type 1 and type 2 are main-tained over their distribution range. We then investigated phenotypic differentiation between type 1A and type 1B, which were found to be so similar that identification based on phenotype is currently not advisable. We propose to treat type 1 and type 2 as distinct species because of their level of genetic and phenotypic divergence, large area of distribution and ample evidence for reduced or absent introgression in contact zones. We maintain type 1A and 1B as subspecies for the time being, pending further analyses of their contact zone. The valid nomen for “Podarcis hispanica type 1 (sensu lato)” is Lacerta muralis guadarramae BOSCÁ, 1916 which becomes Podarcis guadarramae (BOSCÁ, 1916). Lineage type 1A is here described as a new taxon: P. guadarramae lusitanicus ssp. nov., inhabiting northern Portugal and northwestern Spain. The type 1B lineage corresponds to the nominotypical subspecies that inhabits Spain, mostly the Central Iberian Mountains. We were unable to locate an available nomen for “Podarcis hispanica type 2”, which is here described as Podarcis virescens sp. nov. This species is widely distributed in the plains and plateaus of central and parts of south-western Spain as well as central and southern Portugal.

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

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Current climate change-forced local extinctions of ectotherms in their warmer distribution limits have been linked to a reduction in their activity budgets by excess of heat. However, warmer distribution limits of species may be determined by biotic interactions as well. We aimed to understand the role of thermal activity budgets as drivers of the warmer distribution limit of cold-adapted mountain ectotherms, and the colder distribution limit of partially sympatric thermophilous species. In the southern slopes of the Sierra de Guadarrama, Madrid, Spain, (1800–2200 m asl), we collected data from surveys of active individuals, thermal preferences, thermoregulation effectiveness, and activity budgets across 12 different sites exposed to different microclimates and habitats. We assessed how abundances of each species were predicted by activity budgets, restriction time, temperature deviation, habitat covers and date. We found that Iberolacerta cyreni abundances are not predicted by heat-restricted activity time as they were absent or rare in the areas where its activity budgets are broader. Conversely, the abundances of the other lizards were positively predicted by the potential activity time. Habitat preferences and date explained also part of the occurrences of the four species. Our results suggest that realized niches of lizards in the Sierra de Guadarrama are a consequence of niche partitioning by temperature-mediated competitive exclusion. The more thermophilous species were physiologically limited by the reduced chances of being active due to the cold, while the cold-adapted species was abundant in those areas where its potential activity is limited by cold-temperatures, but the thermophilous species cannot inhabit. We provide new insights on the ecological processes affecting the distribution of ectotherm mountain organisms, the assemblage of their communities and how climate change could affect them.

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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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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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We studied the phylogenetic relationships among Greek species of Podarcis (Sauria: Lacertidae) as inferred from partial mtDNA sequences. The results were applied on a molecular clock model as to calculate the time of separation among the populations studied. To tune the molecular clock we considered a geological event on which there is a consensus among geologists, namely the separation of Peloponnese from Crete. This event is considered to have occurred between 5 and 5.5 Mya. Taking in account this date we conclude that P. erhardii and P. muralis shared a common ancestor from 9.5 - 10.5 Mya respectively. Primitive P. erhardii, originating from NW Greece, descended southwards splitting in two main clades: One on the landmass which today we recognize as the Cyclades island group, and a second going further to the south. The two species we recognize today as P. peloponnesiaca in Peloponnisos and P. erhardii in Crete actually originate from this second clade. In other words P. erhardii is paraphyletic with respect to P. peloponnesiaca. Genetic distances among, as well as extant distribution of P. erhardii on Crete and satellite island populations, indicate that at least one splitting, one re – colonization and one extinction events occurred on the island.

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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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Die dem agonistischen Verhaltensrepertoire entlehnten Ausdrucksformen der Paarungsbalz und die Paarung selbst von Podarcis guadarramae lusitanicus im natürlichen Lebensraum (Serra de Montemuro/Nord-Portugal) werden beschrieben. Es besteht weitgehend Übereinstimmung mit dem Paarungsverlauf bei Podarcis hispanicus.

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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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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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Molecular studies suggest that the Iberian wall lizard, Podarcis hispanica, forms a species complex with several monophyletic types. In Central Spain two of these types are spatially not isolated and may interact. Sex pheromones are important for species recognition and, thus, differences between lizards’ types in chemicals used in intraspecific communication could lead to reproductive isolation. Analyses by GC-MS showed that the femoral gland secretions of adult males of different types were different. Males of one type had twelve exclusive compounds, and proportions of some shared compounds differed. This presumably would reflect selection for the persistency and efficiency of chemical signals in different environments; less volatile compounds and with a higher chemical stability being favoured in the type of lizards inhabiting more humid climatic conditions. Differential tongue-flick rates to scents from femoral secretions indicated that males were able to detect and discriminate between males of different types based on chemical cues alone. In contrast, females detected but did not seem able to discriminate between scents of the two types of males. Thus, multiple factors might be simultaneously acting either against or in favor of speciation, leading, on the one hand, to genetic differences between types, but, on the other hand, probably precluding an effective reproductive isolation in areas where both types of lizards may interact.

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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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The genus Karyolysus was originally proposed to accommodate blood parasites of lacertid lizards in Western Europe. However, recent phylogenetic analyses suggested an inconclusive taxonomic position of these parasites of the order Adeleorina based on the available genetic information. Inconsistencies between molecular phylogeny, morphology, and/or life cycles can reflect lack of enough genetic information of the target group. We therefore surveyed 28 localities and collected blood samples from 828 lizards of 23 species including lacertids, skinks, and geckoes in the western Mediterranean, North Africa, and Macaronesia, where species of Karyolysus and other adeleorine parasites have been described. We combined molecular and microscopic methods to analyze the samples, including those from the host type species and the type locality of Karyolysus bicapsulatus. The phylogenetic relationship of these parasites was analyzed based on the 18S rRNA gene and the co-phylogenetic relationship with their vertebrate hosts was reconstructed. We molecularly detected adeleorine parasites in 37.9% of the blood samples and found 22 new parasite haplotypes. A phylogenetic reconstruction with 132 sequences indicated that 20 of the newly detected haplotypes clustered in a well-supported clade with another 18 sequences that included Karyolysus galloti and Karyolysus lacazei. Morphological evidence also supported that K. bicapsulatus clustered in this monophyletic clade. These results supported the taxonomic validity of the genus. In addition, we found some parasite haplotypes that infected different lizard host genera with ancient diverging histories, which suggested that Karyolysus is less host-specific than other blood parasites of lizards in the region. A co-phylogenetic analysis supported this interpretation because no significant co-speciation signal was shown between Karyolysus and lizard hosts.

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Growth rates have complex sources and may determine adult body size in organisms with indeterminate growth. Thus, the interpretation of interpopulation differences in body size along geographical gradients requires the examination of growth and to distinguish between the proximal and ultimate causes of it. Several studies support a link between growth rates and habitat production via climatic effects. Environmental constraints, such as food abundance, may be correlated with climatic conditions and could, in turn, limit growth rate because of limited energy availability. We performed a reciprocal transplant experiment to disentangle the effects of environmental and genetic factors on body size interpopulational divergence in the lizard Podarcis guadarramae along a 500-m elevation gradient with contrasting environmental conditions. Our results showed that the growing environment determined growth rates of juvenile lizards, independently of the population of origin. Hatchlings experiencing the high-altitude growing environment, which had colder and more humid climatic conditions, grew faster than those growing in the low-altitude environment. However, mother’s site of origin did not affect growth. We conclude that the drivers of growth rate differences in P. guadarramae lizards are probably related to between-sites differences in water and food availability.

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In temperate areas, seasonal changes entail a source of environmental variation potentially important for organisms. Temperate ectotherms may be adapted to the seasonal fluctuations in environmental traits. For lizards, behavioural adaptations regarding microhabitat selection could arise to improve thermoregulation during the different seasons. However, little is still known about which traits influence microhabitat selection of lizards and their adaptation to seasonality. Here we used Podarcis guadarramae to study the role of potential intrinsic (body size, sex, age) and environmental traits (air and substrate temperatures, wind speed, and sunlight) in the seasonal changes of body temperatures and microhabitat selection of lizards. We measured body temperatures of lizards in the same habitat during the four seasons and compared the climatic variables of the microhabitats selected by lizards with the mean climatic conditions available in their habitat. Body temperatures were similar for adult males, adult females, and juveniles within each season, being significantly higher in summer than in the other seasons, and in spring than in winter. The same pattern was found regarding substrate and air temperatures of the selected microhabitats. Wind speed and air temperature did not affect body temperatures, while body length was marginally significant and substrate temperatures and season did affect the body temperatures of lizards. Our results during the whole year support the idea that the seasonality could be the most important factor affecting body temperatures of these temperate species. Regarding microhabitat selection, environmental constraints, as environmental temperatures and wind speed, affected the seasonal changes on behavioural thermoregulation of lizards. This effect was similar between sexes and age classes, and was independent of body size. In addition, importance of sunlight exposure of the selected microhabitats (full sun, filtered sun, or shade) also changed between seasons. Hence, environmental constraints were the main forces driving seasonal changes in microhabitat selection.

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

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

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

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Fire is a critical component of terrestrial ecosystems and essential to understand the composition and diversity of communities in fire-prone regions. Nevertheless, fire does not act alone, and other factors such as land-use type and vegetation structure can also operate at different scales, prompting the response to fire of ectotherm communities such as reptiles. We have evaluated the influence of environmental factors on the abundance, diversity, and richness of reptile communities in Serra da Estrela Natural Park (northern Portugal), at both the landscape and micro-habitat scales. In spring 2014 and 2015, 20 transects were surveyed eight times, the vegetation structure and composition were measured, and the extent of land-use types around each transect calculated in this Mediterranean community. At the landscape scale, reptile abundance was higher in natural woodlands and more complex habitats (higher vegetation heterogeneity and plant richness), and reptile evenness was affected by vegetation heterogeneity and time since fire. By contrast, species richness was not related to any environmental factor measured in this study. Only two lizard species were common in the park, Psammodromus algirus and Podarcis guadarramae. Their relative abundances increased with contrasting habitat variables as the former is a ground-dwelling lizard specialist that prefers heterogeneous (unburnt) habitats and the second is a rock lizard that selects open (burnt) stands. At the micro-habitat scale, the probability of reptile presence, reptile abundance, and reptile richness increased with plant richness and decreased with tree cover. This study highlights the importance of several environmental factors, that operate from micro-habitat to landscape scales to understand the response of reptiles to fire.

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

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

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The geographic distributions of two forms of the Iberian wall lizard (Podarcis hispanicus) in Portugal were determined through extensive field surveys. Predictive models of probability of occurrence were developed for both forms of P. hispanicus, based on multivariate logistic regression of environmental variables. On a coarse scale, the best-fit models suggested that the distribution of the north-western form of P. hispanicus can be largely explained by environmental variables such as altitude, mean annual temperature and number of frost days per year. The distribution of the south-western form of P. hispanicus is also explained by altitude and temperature, but the type of climate also appears important. Predicted probabilities of occurrence broadly match the known distributions range ofthe two forms. Where predicted distributions are not confirmed by field surveys, historical and/or interspecific factors may be more important than environmental variables in influencing the distribution of the lizard.

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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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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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Three reptile species were found in Pessegueiro Island (west coast of Portugal): Chalcides bedriagai, Podarcis carbonelli and Blanus cinereus. The first two are confirmations of species already known, but the discovery of B. cinereus is the first record of an insular population in Portugal. The species richness of the island is higher than the expected by its extension. The conservation status of vegetation on the island is poor but the island is nevertheless included in the Natural Park Southwest Alentejo and Costa Vicentina.

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

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

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

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Los ecosistemas insulares ofrecen numerosos ejemplos de adaptación y conductas que son menos frecuentes en poblaciones continentales. En esta nota se documenta la obser-vación de varios adultos de Podarcis hispanica alimentándose de diferentes especies de insectosatraídos por cadáveres en descomposición de rata negra (Rattus rattus). Esta especie es invaso-ra y cuenta con poblaciones muy numerosas en cada una de las islas que forman el archipiéla-go de Cíes, y sus cadáveres pueden servir de forma indirecta como recurso trófico para laspoblaciones insulares de P. hispanica

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