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The aimof this study was to determine the spatial use of the main biophysical elements present in the agro-silvo-pastoral system (trees and shrubs, fallen trunks and branches, rocky outcrops and pilesof stones/walls) by three lizard speciess: Acanthodactylus erythrurus, Podarcis virescensand Psammodromus algirus. However, A. erythruruswas not found in the ten plots studied. For the remaining two species, the hypothesis tests showed that there are differences in the number of individuals when comparing the fiveplots with grazing with the five plots without grazing,within the montado landscape. Where there are rocky outcrops, P. virescens predominates (especially in grazing areas), while Ps. algirusmainly occurs in areas without grazing. At the same time, there are also differences in the elements used as a perch and as a refuge between the two lizards. This type of work helps the creation of management plans by farmers and owners / decision makers, towardsan efficient conservation of reptiles.

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Alto de São Bento is a granite hill near the city of Évora, with floristic, geological and landscape interest, having been declared a núcleo museológico (NMASB). In this study, the local herpetofaunistic diversity was evaluated, the results of which demonstrated the presence of 12 species of amphibians and 11 species of reptiles in the area. Salamandra salamandra and natterjack toad Epidalea calamita predominate in the sampled amphibians; the most common reptiles are the Psammodromus algirus and the gecko Tarentola mauritanica. Such a contribution of diversity seems regionally important, however the NMASB model does not seem efficient in ist conservation, so a redesign into an ecomuseum is suggested.

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O montado, sistema agro-silvo-pastoril de extrema importância para várias espécies de répteis, foi moldado pela ação humana desde tempos ancestrais e a manutenção das suas características seminaturais depende diretamente da continuação da gestão tradicional. A presente investigação teve como principal objetivo a determinação do impacto da gestão do montado, principalmente do pastoreio, no uso do habitat pelas lagartixas Psammodromus algirus, Psammodromus occidentalis e Podarcis virescens. Os resultados atingidos, por meio da realização de testes de hipóteses, apontam para a ausência de impacto dos vários níveis de intensidade de pastoreio no número de indivíduos encontrado em cada cerca, embora, aparentemente, haja uma alteração do comportamento e uso do micro-habitat pelas lagartixas quando expostas às diferentes intensidades. Estas lagartixas mostraram selecionar micro-habitats com uma estrutura simples, sendo que as diferenças entre as espécies se encontram no uso que estas fazem dos elementos que o constituem; Abstract: Habitat Use by Lizards in Agrosilvopastoral Systems of Montado The montado is an extremely important agrosilvopastoral system to several species of reptiles. It has been moulded by the human activity since the primordial times and its characteristics rely directly on the continuity of the traditional ways of management. The present research`s objective has been the measure of the impact inflicted by the management applied on the montado upon the use of the habitat by the species of lizards Psammodromus algirus, Psammodromus occidentalis and Podarcis virescens. The results suggests that there is no impact on the population of the species studied due to the intensity of the grazing, although there has been noticed a change in the behaviour and usage of the micro-habitat by them, when exposed to the different intensity levels. The lizards seem to prefer simple structured micro-habitats being the main difference between the species in the usage that they give to its physical elements.

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Icosahedral nucleo-cytoplasmic large DNA viruses (NCLDV)-like viruses, which form inclusions in the erythrocyte cytoplasm of reptiles, were previously presented as candidates for a new genus of the Iridoviridae family. The present work describes the distribution of infected lizard hosts and ultrastructural characteristics of the viral inclusions of NCLDV-like viruses from Portugal and adjacent locations in Spain. Giemsa-stained blood smears of 235 Lacerta schreiberi from Portugal and Spain, 571 Lacerta monticola from the mountain Serra da Estrela (Portugal), 794 Podarcis hispanica from several localities in Portugal and Spain, and 25 Lacerta dugesii from Madeira Island, were studied. Infection in L. schreiberi was only found in mountain populations, up to 30% in Serra da Estrela and 9–11% elsewhere. It was absent in lizards from lowlands. Prevalence of infection among L. monticola in Serra da Estrela was 10%; infected lizards were found during March to July and October but not in August and September. Infection in P. hispanica was below 3.3%. Only one infected specimen of L. dugesii was identified by light microscopy. Ultrastructural examination of infected samples revealed that the inclusions are virus assembly sites of icosahedral cytoplasmic iridovirus-like virions. Virions from different host species have different ultrastructural features and probably represent different related viruses.

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The author describes a herpetological trip to Portugal organized by a travelling agency specialized in ecological trips. The group consisted ofeighteen people and was especially interested in reptiles and amphibians, though most of the participants were bird-watchers and nature-lovers too. Day by day the author describes the various reptiles and amphibians that were seen. A table is added, in which the num- bers represent the day the respective animals were spotted. The regions that were visited were: Sao Bras {day I), Guadiana river {day 2), Parque Natural da Ria Formosa (day 3), Huelva and Coto de Dofiana in Spain {day 4), Lagoa and the Monchique (day 5), the rocky coast of Carrapateira, Cabo de Sao Vicente (day 6), Almarjao, close to Sao Bras (day 7). Three species of newts and salamanders were seen, eight species of frogs and toads, two species of turt- les, ten species of lizards and five species of snakes. A list was made of the insects, the birds (123 spe- cies) and plants that were seen. The trip was very successful, with regard to the inte- rest of the group

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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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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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For many years lizard thermal ecology studies have relied on the use of contact thermometry to obtain internal body temperature (Tb) of the animals. However, with progressing technology, an interest grew in using new, less invasive methods, such as InfraRed (IR) pyrometry and thermography, to infer Tb of reptiles. Nonetheless few studies have tested the reliability of these new tools. The present study tested the use of IR cameras as a non-invasive tool to infer Tb of lizards, using three differently body-sized lacertid species (Podarcis virescens, Lacerta schreiberi and Timon lepidus). Given the occurrence of regional heterothermy, we pairwise compared thermography readings of six body parts (snout, eye, head, dorsal, hind limb, tail base) to cloacal temperature (measured by a thermometer-associated thermocouple probe) commonly employed to measure Tb in field and lab studies. The results showed moderate to strong correlations (R2 = 0.84 − 0.99) between all body parts and cloacal temperature. However, despite the readings on the tail base showed the strongest correlation in all three species, it was the eye where the absolute values and pattern of temperature change most consistently followed the cloacal measurements. Hence, we concluded that the eye would be the body location whose IR camera readings more closely approximate that of the animal`s internal environment. Alternatively, other body parts can be used, provided that a careful calibration is carried out. We provide guidelines for future research using thermography to infer Tb of lizards.

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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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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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Skeletochronology used for age identification of a Portuguese population of Podarcis hispanica (Steindachner, 1870) (Sauria, Lacertidae). — The age of 92 specimens of Podarcis hispanica, from Cascais (Portugal) are determined by counting the skeletal growth rings in the diaphysis of ihe femur. In the sample studied, four age groups were observed. The results obtained agree with those reported for other lizards such as Podarcis muralis and Lacerta vivipara.

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

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

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To reduce the impact of biological invasions, we need to understand the behavioural mechanisms that enable some species to be successful invaders. Testing differences in behaviour between sympatric congeneric species with different invasive potential is an opportunity to study specific behavioural traits associated with invasion success. Using the invasive Italian wall lizard, Podarcis sicula, and a noninvasive congeneric, the green Iberian wall lizard, Podarcis virescens, which live in sympatry in a location that is novel for P. sicula, we tested their exploratory behaviour, neophobia and boldness: all traits that should promote invasion success. The invasive P. sicula was more exploratory, bold and neophilic than the sympatric native P. virescens. Native lizards had highly repeatable behaviour, whereas in P. sicula boldness was the only behavioural trait that was repeatable. The behavioural traits of the native species, but not the invasive species, were correlated. A lack of correlation between behavioural traits, as well as a lack of repeatability in two of the three behavioural traits, suggests higher levels of behavioural plasticity in P. sicula, which may also explain the success of this lizard during invasions. Our experiment highlights the potential importance of behavioural traits in invasions and provides insight into why P. sicula is such a successful invader.

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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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Herpetological observations in Portugal and southern Spain (Cota Doñana and Sierra del Norte in Andalusia) In the first week of March 2008 a group of sixteen herpetologists visi- ted the Cota Doñana in the south of Portugal and the Sierra del Norte in Andalusia in Spain to look for reptiles and amphibians. The Cota Doñana is a wetland area close to the Mediterranean, with three types of landscapes (dunes, wetlands and shrub lands). The Sierra del Norte is a mountainous area and one of the warmest parts of southern Spain with shrub lands (maquis), low trees and Mediterranean herbs. A lot of species have been found (see table 2), though in relatively low quan- tities.

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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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Populations should adapt to the climate at their respective localities. Here, we examined differences in thermal biology between two populations of Podarcis hispanica lizards from areas with different climates. Lizards from the cold, northern mountains attained lower field body temperatures than lizards form the warm, southern plains. However, the larger body size and darker coloration of northern lizards resulted in slower cooling rates, which may increase efficiency of thermoregulation. Northern populations selected higher temperatures in a thermal gradient, possibly as an adaptation to heat up before conditions changed, as their mountain environment is unpredictable. Finally, lizards from both populations had similar temperature-dependent locomotor performance curves, although southern lizards were relatively faster considering their smaller body size. We suggest that environmental differences may lead to differences in morphology and locally adapted thermal biology in lizards that might allow maximizing thermoregulation in each local climatic conditions

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Chemical signals are important for mate and species recognition. If variation in chemical signals occurs between populations of the same species, these differences could later preclude mating between populations and lead to speciation. In the Iberian wall lizard, Podarcis hispanica, the lipophilic fraction of femoral secretions of males is mainly a mix of steroids and fatty acids. Among steroids, the most abundant compounds are cholesterol and cholesta-5,7-dien-3-ol, which are implicated in intraspecific communication and sexual selection. Interpopulational differences in chemical signals of males, and in response to these chemicals, could contribute to reproductive isolation between populations, which would explain the known genetic differences between these populations. Chemical analyses indicated that five distinct populations of this lizard from Madrid (Central Spain) differed in the proportions of two steroids (cholesterol and cholesta-5,7-dien-3-ol) in femoral secretions. Moreover, lizards discriminated and had high chemosensory responses (i.e., high tongue-flick rates) to these steroids, but showed interpopulational differences. Lizards from populations with cold temperatures and high relative humidity (i.e., northern Madrid) elicited higher responses to these steroids, whereas the converse occurred for lizards from populations occupying dry and hot habitats (i.e., southern Madrid). Interestingly, the magnitude of the chemosensory responses to cholesta-5,7-dien-3-ol in each population mirrored the abundance of this compound in secretions of males of that population. These results suggest that the importance of cholesta-5,7-dien-3-ol in males’ secretions might be lower for lizards from the southern populations. These differences in the relative importance of chemical signals could explain reproductive isolation and cryptic speciation between populations of this lizard.

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The Iberian wall lizard Podarcis hispanica forms part of a species complex with several morphologically and genetically distinct types and populations, which may or may not be reproductively isolated. We analyzed whether female mate choice based on males’ chemical signals may contribute to a current pre-mating reproductive isolation between two distinct populations of P. hispanica from central Spain. We experimentally examined whether females choose to establish territories on areas scent-marked by males of their own population, versus areas marked by males of the other population. Results showed that females did not prefer scent-marks of males from their own population. In contrast, females seemed to attend mostly to among-individual variation in males’ pheromones that did not differ between populations. Finally, to test for strong premating reproductive isolation, we staged intersexual encounters between males and females. The population of origin of males and females did not affect the probability nor the duration of copulations. We suggest that the different environmental conditions in each population might be selecting for different morphologies and different chemical signals of males that maximize efficiency of communication in each environment. However, females in both populations based mate choice on a similar condition-dependent signal of males. Thus, male signals and female mate choice criteria could be precluding premating reproductive isolation between these phenotypically “distinct” populations

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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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Distribution modelling usually makes inferences correlating species presence and environmental variables but does not take biotic relations into account. Alternative approaches based on a mechanistic understanding of biological processes are now being applied. Regarding lacertid lizards, physiological traits such as preferred body temperature (Tp) are well known to correlate with several physiological optima. Much less is known about their water ecology although body temperature and evaporative water loss (Wl) may trade-off. Two saxicolous lacertids, Algyroides marchi and Podarcis hispanica ss are sympatric in the Subbetic Mountains (SE Spain) were they can be found in syntopy. Previous distribution modelling indicates the first species is associated with mountains, low temperatures; high precipitation and forest cover whereas the second one is more generalistic. Here, we perform two ecophysiological tests with both species: a Tp experiment in thermal gradient and a Wl experiment in sealed chambers. Although both species attained similar body temperatures, A. marchi lost more water and more uniformly in time than P. hispanica ss that displayed an apparent response to dehydration. These results suggest that water loss rather temperature is crucial to explain the distribution patterns of A. marchi in relation to P. hispanica ss, the former risking dehydration in dry areas no matter what temperature is. Ecophysiological traits represent a promising tool to build future mechanistic models for (lacertid) lizards. Additionally, the implications for their biogeography and conservation are discussed.

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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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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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This study was carried out in the Iberian-Mediterranean region of Alentejo (Southern Portugal) to discover which ‘montado’ management practices significantly affected reptile variables (abundance, richness). Field surveys on 30 sampling plots allowed us to identify 370 individuals distributed among 12 terrestrial reptile species (six lizards, one amphisbaenid and five colubrid snakes). The lizard Psammodromus algirus was the dominant species (n = 297 individuals). The highest species richness was comprised of five species, whilst two or three species were recorded in two-thirds of the sampling plots. Principal component analysis over management variables accounted for 71.8% of the total variance and subsequently helped to reveal four agrosilvopastoral types. Among them, the highest reptile abundance largely overlapped the cork oak montado, while higher cattle stocking rates were estimated (trampling index) to occur in the holm oak areas, affecting significantly the reptiles. Regression models also showed that both reptile abundance and richness were significantly related to montado areas, where the shrub layer is well preserved and cattle are either absent or present at low stocking rates. Adequate management of montado, preserving the shrub patchwork, is crucial for the conservation of reptiles. Livestock stocking rates should also be maintained at a sustainable level of 0.2–0.4 cattle/ha.

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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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The occurrence of Apicomplexan parasites in Podarcis wall lizards from the Iberian Peninsula and Balearic islands was studied by amplification and sequencing of the 18S rRNA gene. Species from three genera, Hepatozoon, Sarcocystis and Eimeria were detected. The phylogenetic analysis of the 18S rRNA gene provides unexpected insights into the evolutionary history of these parasites. All Hepatozoon specimens were recovered as part of a clade already identified in lizards from North Africa. The Sarcocystis species, detected in Podarcis lilfordi from Cabrera island, in the Balearic Islands, appears related to Sarcocystis gallotiae, known only from endemic Gallotia lizards from the Canary Islands. Based on the lack of snake predators on this island, this parasite form presumably presents an atypical transmission cycle that uses the same host species as both intermediate and final host through cannibalism, like S. gallotiae. Eimeria is reported for the first time from Podarcis lizards. This study shows the power of detecting multiple different Apicomplexa parasites through screening of tail tissue samples and blood drops that are often collected in reptiles for other purposes.

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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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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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In spite of the importance of chemoreception in sexual selection of lizards, only a few studies have examined the composition of chemical signals, and it is unknown whether and how chemicals provide honest information. Chemical signals might be honest if there were a trade-off between sexual advertisement and the immune system. Here, we show that proportions of cholesta-5,7-dien-3-ol in femoral secretions of male Iberian wall lizards (Podarcis hispanica) were related to their T-cell-mediated immune response. Thus, only males with a good immune system may allocate higher amounts of this chemical to signalling. Furthermore, females selected scents of males with higher proportions of cholesta-5,7-dien-3-ol and lower proportions of cholesterol. Thus, females might base their mate choice on the males` quality as indicated by the composition of their chemical signals.

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Sex recognition is based on color signals in many species of lizards. However, olfactory stimuli are also clearly involved, and many species might rely primarily on chemoreception. We aimed to examine whether color pattern or odors, or a combination of both factors, induce the aggressive response of males of the lizard P. hispanicus. We experimentally manipulated the coloration and odor of male P. hispanicus, thereby creating groups with all combinations between coloration and odor of males or females. Using data from staged encounters, we compared the responses of resident males to the experimental groups of manipulated males and their response to unmanipulated individuals (males and females). Responding males reacted significantly more aggressively to intruders with male odors independent of their coloration, whereas intruders with female odors did not elicit aggressive responses but were preferentially courted, irrespective of their actual sex and body coloration. In addition, intruders with female odors elicited a higher number of tongue-flick explorations than male odor ones. Comparisons with unmanipulated male and female intruders agreed with these expectations. Therefore, at least at close range, odoriferous cues seem to be more important than color patterns in sex recognition and intrasexual aggression by male P. hispanicus. We suggest that this might be a pattern commoner than expected in many species of reptiles.

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

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In the western part of Alentejo (Portugal) Podarcis hispanica2 was observed in a Cork Oak (Quercus suber) using passages cre- ated by boring insects as their winter quar- ters.

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Sex recognition is based on colour signals in many species of lizards. However, olfactory stimuli are also clearly involved, and many species might rely more on chemoreception. We aimed to examine whether colour pattern or odours, or both, are used in sex recognition and which cues elicit courtship of females by males of the lizard Podarcis hispanica. We experimentally manipulated the coloration and odour of female P. hispanica, thereby creating groups with all combinations between coloration and odour of males and females. Using data from staged encounters, we compared the responses of resident males to manipulated and unmanipulated individuals (males and females). Responding males reacted signi®cantly more aggressively to female intruders with male odours, independently of their coloration. Nevertheless, coloration seemed to be important in long-distance sex recognition since, in the ®rst minutes, females painted as females received a lower number of aggressive responses. Both colour and odour were important in eliciting male courtship. However, females painted as females and with female odours were preferentially courted. Comparisons with unmanipulated male and female intruders agreed with these expectations. Therefore, at close range, odoriferous cues seem to be more important than colour patterns in sex recognition, but female coloration is also useful at long range to deter the aggressive response of males and to elicit courtship in conjunction with odours.

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

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

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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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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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Exotic animal invasions constitute a major threat to biodiversity worldwide. Our assessment determined the core range occupied by the Italian wall lizard (Podarcis siculus) in the urban area of Parque das Nações (Lisbon, Portugal), where it was accidentally introduced two decades ago. Despite the apparent current scenario of non-expansion, the alien species interferes with the local distribution pattern of the native P. virescens, with both lizard species shown to use the available microhabitats differently. The native P. virescens population displays an overall heterogeneous distribution in the urban matrix. On the other hand, the occupancy of the exotic species clusters within the original area of introduction (garden in front of the Lisbon Oceanarium), as well as in adjacent gardens. There, P. siculus replaced the native P. virescens as the only lizard species present. Apparently, there is no known reason there that prevents the alien lizard from colonizing more available geographic area and expanding.

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I report a cannibalism event in an urban population of Podarcid virescens (Lacertidae). The species is the only lacertid lizard in the site: an artificial wall in Universidad Autónoma de Madrid, where the lizard bask in winter sunny days; mostly adults). Within an ongoing study on the trophic ecology of the species, I found a piece of tail in a fecal pellet (29 scale whorls). The sample was dropped in february2023. The detailed examination of the piece (scales morphology and diameters of the piece of tail) allows to identify it as a distal part of the tail of a young individual. The possibility of the scant provenance from Tarentola mauritanica (Phyllodactylidae), scarcely present in the site, was discarded by morphochemical study of the fecal pellet (work in prep.) The current available data: frequency of the predation event (relatively high at the moment), age/SVL of the victim, and season of occurrence, might preliminarily support the predominant hypothesis on lizards’ cannibalism: young individuals as victims, coincident with low prey availability and when individuals must face winter environmental conditions. If so, cannibalism in this Podarcis virescens population (not necessarily in other species) might be important on population structure (agespatial segregation), and population dynamics.

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

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

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

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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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Invasive species can carry parasites to introduced locations, which may be key to understand the success or failure of species establishment and the invasive potential of introduced species. We compared the prevalence and infection levels of haemogregarine blood parasites between two sympatric congeneric species in Lisbon, Portugal: the invasive Italian wall lizard (Podarcis siculus) and the native green Iberian wall lizard (Podarcis virescens). The two species had significant differences in their infection levels: while P. virescens had high prevalence of infection (69.0%), only one individual of P. siculus was infected (3.7%), and while P. virescens exhibited an average intensity of 1.36%, the infected P. siculus individual had an infection rate of only 0.04%. Genetic analyses of 18S rRNA identified two different haemogregarine haplotypes in P. virescens. Due to the low levels of infection, we were not able to amplify parasite DNA from the infected P. siculus indi vidual, although it was morphologically similar to those found in P. virescens. Since other studies also reported low levels of parasites in P. siculus, we hypothesize that this general lack of parasites could be one of the factors contributing to its competitive advantage over native lizard species and introduction success.

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

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

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

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The author joined a herpetological journey ro Extremadura in Spain. The group consisred of eigllteen per- sons and was travelling vith two vans. The area is more or less as large as the Nerherlands, but Jess popula- ted. The largestpart ofthis region is occupied by `dehesa`, an open mediterrenean wood1311dscenery with catrle-breeding. Moreover there is steppe coumry, mounrains in the north (Sierra de Gredos) and finally hilly cotumy (Sierra de Guadalupe a11d Monfragüe). Day by day the areas that were visited a11d rhe reptiles and 3lnphibians that were seen are described. In total twelve species of a111phibians and twenty species of reptiles are found.

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Eine Reise nach Portugal führte mich mit befreundeten Eidechsenhaltern in die Serra de Sấo Mamede. Vom 22.04. bis 28.04.2017 bereisten wir den 31.750 Hektar großen Nationalpark, um nach Reptilien, insbesondere nach Eidechsen, zu suchen. In der Literatur werden bei den Schlangen acht Arten und bei den Eidechsen zehn Arten aufgelistet. Davon sind sechs Arten den Halsbandeidechsen zugeordnet. Ein Großteil der in Portugal vorkommenden Reptilien war also in dem kleinen Gebiet zu erwarten.

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Die Autoren berichten, wie sie bei zwei unfteiwilligen Zwischenstopps in Madrid auf eine für sie neue Eidechse stießen, die vier Jahre später dann sogar als neue Art beschrieben wurde.