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Las restricciones térmicas pueden limitar la fisiología y el comportamiento de los ectotermos debido a la dependencia térmica de sus funciones metabólicas. Los animales termorreguladores han desarrollado mecanismos de ajuste, tanto comportamentales como fisiológicos, que permiten mantener sus temperaturas corporales en el rango de temperaturas óptimas para la mayoría de las funciones fisiológicas. Además, ciertos rasgos morfológicos pueden favorecer el intercambio de energía con el ambiente así, un menor tamaño corporal aumenta las tasas de calentamiento, del mismo modo que los colores oscuros, por su mayor capacidad de captación de la radiación. En esta tesis, se analiza la termorregulación globalmente, manejando varias especies de saurios para dilucidar los mecanismos que optimizan el proceso de termorregulación en los ambientes más desfavorables. Utilizando como modelo una especie de alta montaña, Iberolacerta cyreni, se estudian los mecanismos de termorregulación empleados en ambientes óptimos y sub-óptimos, explorando el papel del tamaño corporal como rasgo morfológico determinante en la biología térmica. En segundo lugar, se establece el efecto de la temperatura sobre el crecimiento juvenil temprano, rasgo relevante por su influencia sobre el tamaño corporal en la madurez, comparando dos poblaciones de ambientes térmicos contrastados en Iberolacerta monticola. El estudio del rol del melanismo en la biología térmica se aborda desde dos perspectivas. Primero, utilizando como modelo la especie I. monticola, se comparan los patrones de coloración en dos poblaciones de los extremos de su distribución altitudinal, analizando simultáneamente las diferencias locales en la cantidad de melanina y comparando los valores actuales con los de especímenes preservados recolectados en las mismas poblaciones en 1983-1984, discutiendo los posibles efectos del cambio climático. Además, se determina el papel del melanismo sobre las tasas de calentamiento bajo distintas condiciones térmicas. Por último, se midió la variación en la reflectancia y el tamaño corporal, así como en respuesta térmica en la especie Cordylus cordylus, endémica de Sudáfrica. Analizando los resultados bajo la hipótesis del melanismo térmico, se evalúa la hipótesis de la co-adaptación entre la fisiología térmica, el tamaño corporal y el melanismo, valorando la influencia de otros factores de selección, como la cripsis o la disponibilidad de refugios adecuados. Nuestros resultados indican que I. cyreni es una especie termorreguladora activa bastante precisa pese a las limitaciones de las temperaturas disponibles en el campo, mostrando temperaturas preferidas inferiores a la mayoría de lacértidos. El mantenimiento de la temperatura corporal se realiza fundamentalmente mediante ajustes comportamentales, incluso en ambientes sub-óptimos con elevados costes asociados. Por otro lado, en I. monticola, existen diferencias locales en el crecimiento temprano en masa independientes del ambiente térmico, mientras que la norma de reacción del crecimiento en longitud mostró patrones opuestos entre poblaciones. Además, se encontraron evidencias de crecimiento compensatorio, con patrones opuestos en la interacción entre la población y el ambiente térmico entre las poblaciones. El grado de melanismo en I. monticola varía dependiendo de la población (altitud), el sexo y el periodo temporal (1983-1984 vs 2011-2012), con lagartijas más oscuras a mayor altitud, machos más oscuros que hembras, y mayor dimorfismo sexual en la población de montaña en la muestra actual (2011-2012). Este patrón complejo de variación sugiere la existencia de diferentes procesos y mecanismos causales que actúan sobre la evolución de la coloración. Por último, los resultados muestran que en C. cordylus, el tamaño corporal, el melanismo y las adaptaciones térmicas podrían estar co-adaptadas, aunque otros factores, como la selección sexual y la cripsis podrían intervenir en la variación del melanismo entre poblaciones.

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En Castilla y León, en 2021, la Junta de Castilla y León determina el comienzo del proyecto `Estudio para el seguimiento del estado de conservación de los anfibios y reptiles de Castilla y León`. Se realizó el muestreo dirigido sobre 18 cuadriculas UTM 10x10 km (2 por provincia) seleccionadas sobre hábitats favorables de las 13 ZEC propuestas en el Plan de Monitorización del Estado de Conservación de la Biodiversidad; de los Espacios Protegidos de la Red Natura 2000 y de los Espacios Naturales Protegidos de Castilla y León. La metodología es tomada del programa SARE (Seguimiento de los Anfibios y Reptiles Españoles). Además, dentro de los objetivos, se llevó a cabo la revisión bibliográfica y el muestreo específico para aumentar el conocimiento de tres especies de anfibios de interés comunitario: R. dalmatina, C. lusitanica e I. alpestris. Por un lado, la sequia del año 2022 ha producido malos resultados en los muestreos de anfibios con lo que habría que repetir el esfuerzo realizado sobre las especies prioritarias y por otro se han obtenido abundancias más bajas que el año anterior. En anfibios la especie que alcanzó mayor densidad en la mayoría de las estaciones donde estaba presente resultó ser E. calamita, seguido de L. boscai y P. perezi,. La media de la densidad máxima de anfibios ya metamorfoseados es mayor para los anuros que para los urodelos. La densidad larvaria máxima no varió significativamente por regiones biogeográficas, aunque los anuros fueron ligeramente más abundantes en la región mediterránea que en la atlántica. Las densidades larvarias variaron entre especies, siendo especialmente abundantes en A. obstetricans, P. cultripes, R. parvipalmata y T. pygmeus. Las especies de reptiles más abundantes durante los dos años de muestreo fueron: M. leprosa, I. cyreni, I. martinezricai, E. orbicularis, P. algirus, I. monticola y P. muralis. La densidad máxima varió significativamente entre grupos taxonómicos, con una densidad alta en quelonios, media en saurios, y muy pequeña en ofidios. Para calcular tendencias poblacionales se requieren tres años de seguimiento, hasta que no termine el 2023, no podemos valorar una posible evolución. No obstante hemos obtenido conclusiones metodológicas. Por ejemplo la inclusión de estaciones de muestreo especialmente favorables en zonas limítrofes a las cuadrículas UTM 10x10 ha sido acertada y, de hecho, probablemente hubiera sido todavía más acertado elegir las estaciones de muestreo sin atender a los límites de las cuadriculas. Los resultados corroboran que el área de estudio presenta poblaciones abundantes de anfibios y reptiles, aunque no exentas de problemas y muy variables en importancia según la zona geográfica, destacando la riqueza de las poblaciones de quelonios. También, y como cabría esperar, la biología particular de ofidios y quelonios aconseja la realización de censos específicos para estos grupos con objeto de obtener medidas más realistas de sus abundancias.

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The genetic structure and relationships of five populations of the Iberian group of Iberolacerta (sensuArribas 1999) were analysed by protein electrophoresis. In this study we confirmed the differentiation between the group of populations from Galicia/Cantabrian/S. Estrela versus the group of populations from de Spanish Central System, Gredos and Guadarrama that are included in the two different species by Arribas (Herpetozoa 9(1/2), 31–56, 1996; Russian J. Herpetol. 6, 1–22, 1999), Iberolacerta (I.) monticola and Iberolacerta (I.) cyreni, respectively. However, the differentiation level is not high enough to clearly prove their specific discrimination. On the other hand, we did not confirm the subspecific discrimination of the Gredos –Iberolacerta (I.) cyreni castiliana, and Guadarrama –Iberolacerta (I.) cyreni cyreni populations, proposed by Arribas (1996). These two populations are genetically almost homogeneous. Interestingly, we found an unexpected high genetic similarity between the Galician and the Serra da Estrela populations, presently included into two different subspecies, Iberolacerta (I.) monticola cantabrica and Iberolacerta (I.) monticola monticola, respectively. Their genetic similarity is even higher than that between the populations of Galicia and the Cantabrian Mountains, which are both included in the same subspecies, cantabrica. This result suggests that the populations of Galicia and Serra da Estrela would have maintained contacts, possibly through the north of Portugal, until relatively recent times. Their separation is thus probably post-glacial. Some evidence also points to the existence of relatively recent contacts between the population of Serra da Estrela and those of the Central System, particularly, with the neighbouring Peña de Francia population. With the cautions imposed by the reduced sample size of our analysis, the significant differentiation of the populations from Galicia and the Cantabrian Mountains allow us to suggest that this last population may not be the result of a recent expansion of the Galician population as Arribas (1996) suggests, but, more likely, the result of a fragmentation process of a more ancient and wider north-eastern distribution area of this group of rock-lizards.

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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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Lizard erythrocytic viruses ~LEVs! have previously been described in Lacerta monticola from Serra da Estrela, Portugal. Like other known erythrocytic viruses of heterothermic vertebrates, these viruses have never been adapted to cell cultures and remain uncharacterized at the molecular level. In this study, we made attempts to adapt the virus to cell cultures that resulted instead in the isolation of a previously undetected Ranavirus closely related to FV3. The Ranavirus was subsequently detected by polymerase chain reaction (PCR) in the blood of infected lizards using primers for a conserved portion of the Ranavirus major capsid protein gene. Electron microscopic study of the new Ranavirus disclosed, among other features, the presence of intranuclear viruses that may be related to an unrecognized intranuclear morphogenetic process. Attempts to detect by PCR a portion of the DNA polymerase gene of the LEV in infected lizard blood were successful. The recovered sequence had 65.2/69.4% nt/aa% homology with a previously detected sequence from a snake erythrocytic virus from Florida, which is ultrastructurally different from the studied LEV. These results further support the hypothesis that erythrocytic viruses are related to one another and may represent a new group of nucleocytoplasmic large deoxyriboviruses.

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The study of the effect of parasites on their host populations is essential for understanding their role in host population dynamics and ecology. We describe the general field population biology of haemogregarines in the Iberian rock lizard, Lacerta monticola, examining the factors that determine the prevalence and intensity of infection. Prevalence and infection intensity were higher in adults than in juvenile lizards. The prevalence rate was higher in larger lizards, probably because they were also older and had been more often in contact with parasites during their lifetime. During the mating season, the intensity of parasite infection was greater in males than in females, probably due to immunosuppressive effects of testosterone. The parasite load had a negative effect on the body condition during the reproductive season. The results suggest that the interactions between parasites and hosts are not stable in this lizard population.

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Previously known in Portugal only in Serra da Estrela (Centre of Portugal), the Iberian rock-li zard, Lacerta monticola, is now reported inside the Montesinho Natural Park (North of Portugal). The specimens found in Museu Bocage`s collection could be tentatively assigned to subspecies call1abrica.

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

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We conducted a field study to examine the significance of the seasonal changes in activity levels, spatial relationships, and social interactions of the Iberian rock lizard, Lacerta monticola, during the same favorable climatic period. Activity level, movement rate, home-range size, number of same-sex neighbors, and number of agonistic interactions decreased as the mating season finished for males but not for females. The number of females` home ranges overlapped by the home range of each male was also higher in the mating season than in the nonmating season. Measurements of available ambient temperatures showed that the thermal environment was suitable for the activity of lizards in both seasons, therefore the seasonal changes cannot be explained by thermal constraints. Males gain access to several females by increasing their home-range size, thus overlapping a larger number of females` home ranges, but as a consequence, also those of a larger number of male competitors. Thus, the cost of incurring agonistic interactions also increases. These results suggest that because activity and aggressiveness in males are related to their mating success, the balance between the costs and benefits of their activity and spatial strategies differs seasonally, and this may be the cause of the observed seasonal changes.

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The ability of territorial lizards to discriminate between scents of neighbors and non-neighbors might contribute to decreasing the costs of aggressive interactions. To test this hypothesis, we conducted a field study to analyze the spatial relationships between male Iberian rock-lizards, Lacerta monticola. We then used the same individuals in a laboratory experiment to test whether male lizards can use chemical cues to discriminate between familiar conspecific males (those whose home ranges overlapped) and unfamiliar conspecific males (those whose home ranges did not overlap, and whose home range centers were at least 50 m apart). Differences in tongue-flick rates in the presence of chemical cues suggested that male L. monticola discriminated between odors of familiar and unfamiliar males. The behavioral responses were also dependent on relative differences in body size between the responding male and the unfamiliar male that donated the scent: There was a significant negative correlation between tongue-flick rates emitted in cages of unfamiliar males and the body size differences between males. In contrast, when the donor of the scent was a familiar male, the tongue-flick rate was not dependent on body size differences. These results are compatible with individual discrimination through chemical cues in male L. monticola.

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We conducted a field study to analyze spatial relationships among male Iberian Rock Lizards, Lacerta monticola. We then used the same individuals in a laboratory experiment to test whether avoidance responses of a male lizard in a previously unknown area is affected by presence of scents of familiar and unfamiliar resident males. Time spent attempting to escape in presence of unfamiliar odors was significantly higher than in presence of familiar odors suggesting avoidance of unfamiliar scent marks, which may lower the costs of aggressive interactions. Our results are concordant with previous studies in which differential tongue-flick rates showed discrimination between odors of familiar and unfamiliar males of L. monticola.

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Many studies on contest competition used residency asymmetry as a discrete variable. However, the probability of winning an interaction may change as a continuous function of the value of the location where the encounter occurs. We performed a field study to examine the importance of location within a home range and relative body size to the outcomes of agonistic interactions between male lizards, Lacerta monticola. The distances to activity centers (themost used locations based on a density function of sightings) and relative size play important roles in agonistic interactions and had interacting effects in natural conditions. On the other hand, previous studies with lizards suggested that inferior competitors are able to avoid agonistic interactions in the field. Thus, we staged encounters in the laboratory to examine the behavioral responses of smaller individuals. The responses of each focal smaller male were measured in its own home cage (resident), in the cage of a larger male (intruder) and in a cage in which no male was previously present (control). The predominant behavioral tactics of smaller males were avoidance when they are the intruders and displaying when they are the residents. Submissive displays by smaller males may help reduce the costs of agonistic encounters.

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Theoretical and field studies suggest that inferior competitors may be able to avoid costly agonistic interactions, although the mechanisms involved are poorly understood in lizards. The “Dear enemy recognition” theory proposes that higher tolerance among familiar than among unfamiliar conspecifics may reduce costly interactions. Male lizards Lacerta monticola are able to discriminate between familiar and unfamiliar conspecific males through chemical signals alone. To better understand the implications of this discrimination ability, we performed a laboratory experiment on the role of familiarity in social tolerance estimated as the degree of spatial proximity with no sign of aggressiveness between males with a size asymmetry. In natural conditions the outcome of neighbour-stranger interactions might be the result of asymmetries in residence. Therefore, we staged pairwise encounters between familiar and unfamiliar males in neutral terraria to avoid an interacting effect of residency asymmetry. The latency to the first time in close proximity was significantly longer when small males were faced with unfamiliar males. The time that unfamiliar lizards spent in close proximity was significantly lower than that with familiar males. Our design allowed us to conclude that there was a higher social tolerance among familiar individuals with a size asymmetry that was independent of residency asymmetry.

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Theoretical and field studies suggest that inferior competitors may be able to avoid costly agonistic interactions, although the mechanisms involved are poorly understood in lizards. The “Dear enemy recognition” theory proposes that higher tolerance among familiar than among unfamiliar conspecifics may reduce costly interactions. Male lizards Lacerta monticola are able to discriminate between familiar and unfamiliar conspecific males through chemical signals alone. To better understand the implications of this discrimination ability, we performed a laboratory experiment on the role of familiarity in social tolerance estimated as the degree of spatial proximity with no sign of aggressiveness between males with a size asymmetry. In natural conditions the outcome of neighbour-stranger interactions might be the result of asymmetries in residence. Therefore, we staged pairwise encounters between familiar and unfamiliar males in neutral terraria to avoid an interacting effect of residency asymmetry. The latency to the first time in close proximity was significantly longer when small males were faced with unfamiliar males. The time that unfamiliar lizards spent in close proximity was significantly lower than that with familiar males. Our design allowed us to conclude that there was a higher social tolerance among familiar individuals with a size asymmetry that was independent of residency asymmetry.

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Activity, temperature and habitat selection of Lacerta monticola were studies at one locality in the Cantabrian mountains. L. monticola has an unimodal pattern and is relatively inactive after midday. There is no significant different in habitat use between males, females and subadults. Lizards do not seasonally vary their patterns of habitat utilization. There is a significant correlation between body temperature and ambient temperature. Also, body temperature is significantly correlated with substrate temperature. Males, females and subadults did not differ significantly in body temperature. Body temperatures varies with time of day but not among months.

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The ability to shed (autotomize) all or part of the tail, usually in response to predator attack, and often to subsequently regenerate it is widespread in lizards and amphisbaenians and also occurs in a few snakes and in the tuatara. Most species possess a sophisticated intravertebral autotomy mechanism which seems to be primitive in the Squamata. This appears to have been independently lost in members of many groups, but some agamids and snakes have regained the ability to shed their tails by a simpler intervertebral means and a -number of agamids have also redeveloped tail regeneration as well. Breakable tails are used to evade capture in two main ways: by enabling reptiles to break away from predators that have grasped them by the tail and by providing a distraction which deflects the attention of the attacker away from the vulnerable head and body. It is argued that loss of caudal autotomy has occurred when the costs of tail shedding outweigh its benefits. Likely costs include the expense of regrowing the tail and the loss of a variety of possible tail functions that may cause partial incapacitation, at least until the tail regenerates. Benefits of autotomy are liable to be low if predation is rare, if the animal is able to protect itself effectively in other ways, if it is too slow to evade further pursuit after the tail is shed, or if the tail is small or unpalatable and consequently not likely to distract a predator. Benefit variation may well be greater than cost variation and therefore more important in initiating the loss of autotomy mechanisms. Many taxa that do not shed the tail appear to conform to the above interpretation, but in some cases, such as the Platynota, Agamidae and Chamaeleonidae, lack of intravertebral autotomy may reflect the history of these groups rather than being a direct result of present ecological pressures. The distribution of intervertebral autotomy in the Agamidae suggests that it may have evolved only in rather special circumstances where tail fragility is advantageous even in the absence of the ability to regenerate. Restriction ot autotomy planes to the tail-base, so that the whole organ is lost, a condition found in a number of relatively slow-moving geckoes, is interpreted as a means of ensuring that enough of the tail is shed to distract a predator from further pursuit. The stimulus necessary to induce autotomy can vary rapidly in individual lizards and at least some of these changes probably maximize the effectiveness of the tail-shedding mechanism. Differences in the readiness with which all or part of the tail is shed exist between species and are likely to reflect the balance of costs and benefits in particular cases. Variations in incidence of broken tails between species and populations may be due to such differences in fragility but many other factors may play a part, including the age structure of samples, incidence of unsuccessful attacks by predators and ability to evade predators after autotomy. There is a clear tendency for climbing lizards, especially those living on rock surfaces, to have higher incidences of broken tails than ground-dwelling species, perhaps because the tail is usually less important in locomotion in the first group. Many lizards possess conspicuously coloured tails and tail movements that seem likely to help distract attention from the head and body. Conspicuous tail colouring is more frequent and often better developed in young animals, which tend to be more vulnerable than adults, and in active species from open habitats where crypsis may not always be very effective. Conspicuous tails usually have contrasting light and dark areas in nocturnal forms but are often a single bright colour in diurnal ones, probably reflecting the visual capacities of their respective predators. The predominance of blue tails in day-active species may be because this colour is striking close to but not very arresting at a distance, so it may not attract predators from far away while still drawing their attention at close quarters.

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

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DNA sequence indicates the Lacertidae contain two subfamilies, Gallotiinae and Lacertinae, the latter comprising two monophyletic tribes, the Eremiadini of Africa and arid southwest and central Asia, and the Lacertini of Europe, northwest Africa and southwest and east Asia. Relationships within the 108 species of Lacertini are explored using mtDNA (291 bp cytochrome b; 329 bp 12S rRNA for 59 nominal species, and reanalysis of the data of Harris et al. 1998, and Fu 2000). The morphology of the tribe is reviewed and 64 of its characters (equivalent to 83 binary ones) also used to assess relationships. The Lacertini are assigned to 19 monophyletic units of 1 to 27 species, recognised here as the following genera (contents are indicated in brackets): Algyroides, Anatololacerta gen. nov. (L. danfordi group), Apathya (L. cappadocica group), Archaeolacerta (L. bedriagae), Dalmatolacerta gen. nov. (L. oxycephala), Darevskia (L. saxicola group), Dinarolacerta gen. nov. (L. mosorensis), Hellenolacerta gen. nov. (L. graeca), Iberolacerta (L. monticola group), Iranolacerta gen. nov. (L. brandtii and L. zagrosica), Lacerta s. str. (sand and green lizards, L. agilis group), Parvilacerta gen. nov. (L. parva and L. fraasii), Phoenicolacerta gen. nov. (L. laevis group), Podarcis (wall lizards), Scelarcis (L. perspicillata), Takydromus (Asian grass lizards), Teira (L. dugesii), Timon (ocellated lizards, L. lepida group) and Zootoca (L. vivipara). Both mtDNA and morphology indicate that Lacerta and Timon are sister taxa, and DNA suggests further possible relationships among genera (Fig. 1, p. 6). Neither DNA nor morphology indicates that the archaeolacertas (sometimes formalised as Archaeolacerta sens. lat.) form a clade. Instead, they are representatives of an ecomorph associated with living on rock exposures and using the narrow crevices that these contain. The Lacertidae probably arose in the European area, with the Gallotiinae later reaching Northwest Africa and the Canary Islands, and the ancestor of the Eremiadini invading Africa in the mid-Miocene. The Lacertini spread through much of their present European range and diversified, perhaps largely by repeated vicariance, around 12–16 My ago, producing the ancestors of the present mainly small-bodied genera, which then underwent often modest speciation. Three units spread more widely: the Lacerta-Timon clade of large-bodied lizards probably dispersed earliest, followed by Algyroides and then Podarcis. Overall, European Lacertidae show a pattern of repeated spread, often accompanied by restriction of previous groups. Expansion of Lacertini may have displaced earlier lacertid lineages from all or much of Europe; while spread of Podarcis may have restricted many other genera of Lacertini. The earlier expansion of the Lacerta-Timon clade probably did not have this effect, as difference in adult body size restricted competitive interaction with other forms. Several invasions of more distant areas also occurred: of East Asia by Takydromus over 10 My ago, and more recently of northwest Africa by Podarcis, Scelarcis and Timon, and Madeira by Teira. Relationships within the Eremiadini estimated from both mtDNA, and nDNA differ considerably from those based on morphology. They indicate relatively mesic forms may have diversified widely across Africa and given rise to at least three independent invasions of arid habitats. MtDNA also indicates that Lacerta andreanskyi belongs in the Eremiadini and may occupy a basal position there. It is assigned to a further new genus, Atlantolacerta gen. nov.

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The present morphological study analyzes the geographical variation of `Lacerta` monticola BOULENGER, 1905 and `L..` cyreni MÜLLER & HELLMICH, 1937 by means of univariate and multivariate statistical analyses. `L. ` monticola shows a low degree of morphological interpopulation differentiation (considerable overlap of the ranges of respective character states). Yet, one sample (Serra da Estrella) is separable from the rest of the Cantabric samples. The first corresponds to `L.` m. monticola BOULENGER, 1905, the second to `L.` m. cantabrica MERTENS, 1929. `L. ` cyreni forms several allopatric populations which have reached a considerable degree of divergence. The populations of Gredos seem to be subspecifically distinct from the Guadarrama ones `L. ` c. cyreni MÜLLER & HELLMICH, 1937) and the small endangered Pena de Francia population is strongly different from both.

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The hemipenis of all the Iberolacerta species is studied. Overall morphology is very similar among the different species. Hemipenes are symmetrically bilobed, with the apical section shorter than the basal one (I. galani, I. cyreni, I. monticola monticola, I. horvathi, I. bonnali, I. aranica and I. aurelioi) or subequal or with slightly inverted proportions (I. m. astur and I. martinezricai). All are similarly sized and proportional to the animal’s SVL, except in I. aurelioi (and perhaps to a lesser extent in I. horvathi) that have notably small hemipenes. Concerning the hemipenial microornamentation, all the Iberolacerta s. str. (I. monticola, I. galani, I. martinezricai, I. cyreni and I. horvathi) have hemipenial microornamen-tation of crown-shaped tubercles, more or less elongated with a series or a crown of small spinules in their apex. Among these, isolate odd shaped tubercles can appear, probably corresponding to immature states that do not reach their complete development. The Pyrenean species (subgenus Pyrenesaura) are variable. Iberolacerta bonnali and I. aranica have mainly spiny-like tubercles (among which some crown shaped can appear), whereas I. aurelioi presents crown-shaped microornamentation (with scattered spiny-like, as occur in other Iberolacerta species). Our hypothesis is that crown shaped models are derived (or more mature, following an ontogenetic criterion) in respect to the spiniform ones, but some spiny (or finger-shaped) can be secondary reversals by incomplete maturation. These secondary derived and the original primitive spiny-shaped ones are very difficult if not impossible to distinguish. Phenomena as the heterochronies could be beneath these reversals in maturation.

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The microornamentation of the scales (oberhautchen) has been studied by means of enamel casts (in negative), or with detached skin pieces (positive cast in the upper face or negative in their lower face of the scales) and its observation by transparency with optical microscopy at 40x, 400x and 1000x. In this study we have found a remarkable uniformity in all genera of Lacertini studied (already known, although significantly expanding the number of species previously studied). In light of the results obtained, it seems that the use of the microornamentation of the scales in taxonomy is limited. There is a remarkable degree of variation and probably of homoplasy, as it happens in so many characters of Lacertini. Nor is it clearly linked to certain conditions of the environment, although it may have a certain importance in mechanisms such as evapotranspiration and perhaps in the crypsis through its influence on the reflectance of the scales. These scales can present protuberances (like small tubercles) on their surface (derived character) in Algyroides (the four species), and in the case of Algyroides marchi also seems to present small pits on its scale surface. In most lacertids the cells of the “oberhautchen” are usually narrow (or short anteroposteriorly, if preferred), between 1 and 4 μm, having the appearance of ribbons or transverse straps. Smaller sizes (derivated) appear more or less marked in Algyroides spp, Darevskia praticola complex, Parvilacerta spp and in Dinarolacerta mosorensis (although not in D. montenegrina) and is difficult to interpret. Except in Dinarolacerta, in the other species it coincides that they are ground or ground-rock interface species, and the coarser stripes could have to do with reflectance (imitating the vegetation or the medium between which they move, according to the hypothesis of Allam et al.). In the case of Dinarolacerta, very saxicolous, it is probably the sister group of Algyroides and the common morphology could have a phylogenetic origin linked to its relationship with Algyroides. There is also no relationship with moisture since some come from dry places and others are from very wet areas. The posterior flange of the cells may be slightly mounted in the next cell (primitive character) but appears quite raised (derived character) in Algyroides nigropunctatus (much less in A. moreoticus and A. marchi, and apparently little or nothing in A. fitzingeri). The rear flange of the cells can be smooth, corrugated or notched. In our samples, irregular (and even dentate) posterior ridges appear in numerous Darevskia such as D. derjugini, D. parvula, D. raddei complex [D. raddei raddei, D. r. vanensis and D. nairensis], in part of the group of D. rudis [D. rudis sspp. and D. bithynica sspp.], several Iberolacerta (I. bonnali, I. horvathi and I. monticola astur), as well as in Teira dugesii. The character appears, although less marked in Algyroides (more in A. moreoticus, but less marked or barely visible in the others), in Hellenolacerta graeca, and other Iberolacerta (I. aurelioi, I. martinezricai, and scarcely marked in I. cyreni). Very little marked in Darevskia mixta, somewhat more in D. dryada and none in D. clarkorum, and also appears in the group of D. saxicola (D. saxicola, D. brauneri brauneri, D. b. darevskii, D. lindholdmi, D. szczerbaki), Dalmatolacerta oxycephala, Dinarolacerta mosorensis (only in the Biokovo sample, and not in Durmitor or D. montenegrina), Apathya cappadocica, in some places of the scales of Parvilacerta fraasii (but not in P. parva) and very little in D. (p) hungarica; also in the remaining group of D. rudis (D. valentini valentini, D. v. lantzicyreni, D. rudis bolkardaghica, D. portschinskii), and some parthenogenetic species (D. armeniaca, D. dahli, D. rostombekowi) but not in D. unisexualis, D. uzzelli, D. sapphirina, D. bendimahiensis or in the triploid hybrid between D. unisexualis and D. nairensis. In these parthenogenetic species it seems a general rule that the character of the crenulation or irregularity of the posterior flange of the cells resembles that of the paternal rather than the maternal species. The exception is the triploid hybrid [arisen from the hybridization of D. unisexualis (parthenogenetic female, without the character) and a male of D. nairensis (with fairly developed character)] that lacks character, and in this case is more similar to the maternal species of which has two genetic sets compared to only one of the paternal one.

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Iberolacerta populations from the Northern Montes de León (NML) were studied by means of external morphology (scalation and biometry), osteology and genetics (mtDNA and microsatellites), searching for their homogeneity (“intrazonal analysis”) and, once verified, comparing them with Iberolacerta monticola s. str. (from Central Cantabrian Mountains) and I. galani (from Southern Montes de León) (“extrazonal analysis”) from neighboring areas. Our “intrazonal analysis” revealed discordances between the different approaches, especially the patterns of variation of nuclear microsatellites (congruent with external morphology) and mtDNA, namely a very low nuclear differentiation between relatively highly differentiated mtDNA lineages. The morphological approach was unable to discriminate any of the populations as significantly different from the others in the NML. Mitochondrial DNA revealed a haplotype lineage closely related to I. galani (MNL-II in our text) in some specimens of Sierra de Villabandín and Suspirón, but these populations are morphologically indistinguishable from the main part of the other populations that belong to lineage NML-I, phylogenetically closer to I. monticola. After a separation from I. monticola ca. 1.8 Mya, the populations in this geographic region must have suffered at least two different waves of gene flow from I. galani, the second one not much later than 0.5 Mya. Microsatellite results indicate that all the NML populations are genetically similar in terms of their nuclear genomes, independently of their mitochondrial differentiation (NML-I vs. NML-II haplotype groups). Since all the morphological and microsatellite evidences point towards the fact that, independently of the mitochondrial haplotypes that they bear (NML-I or NML-II), there is only one taxon in the area, we describe it as: Iberolacerta monticola astur ssp. nov. Concerning the relationships of I. m. astur ssp. nov. with I. monticola s. str. and I. galani (“extrazonal analysis”), in the female analyses the new taxon centroid is closer to I. monticola s. str. than to I. galani (more similarity with I. monticola s. str.), whereas in the male analyses the relationship is just the contrary (closer to I. galani, paralleling the direction of the hypothesized past hybridization). Moreover, in both sexes’ ANOVA, I. m. astur ssp. nov. results more similar (less P<0.05 differences) to I. galani than to I. monticola s. str. Osteologically, I. m. astur ssp. nov. is slightly more similar to I. monticola s. str. than to I. galani, especially in the squamosal bone, which is regularly arched (primitive shape). Genetically, as indicated above, the NML populations can be subdivided in two groups according to their mitochondrial DNA, namely NML-I (bearing clearly differentiated haplotypes, phylogenetically closer to I. monticola) and NML-II (whose haplotypes could have been mistaken for those of an I. galani population). This mitochondrial subdivision has at most a subtle nuclear correlate, however. According to the nuclear microsatellite markers, all the NML populations belong to a single group (I. m. astur ssp. nov.), which would be more similar to I. galani than to I monticola, with NML-II populations lying closer to I. galani than those from the NML-I group and, correspondingly, more distant from I. monticola. The discordant phylogenetic signal of mitochondrial and nuclear markers is discussed in terms of past introgression events and sex-biases in phylopatry and dispersion in these species. Iberolacerta monticola astur ssp. nov., inhabits the Northern Montes de León (Sierra de Gistreo sensu latissimo): Gistredo, Catoute, Tambarón, Nevadín, Villabandín (or Macizo del Alto de la Cañada), Arcos del Agua (or Fernán Pérez), Tiendas and Suspirón, mainly in quartzite and slate rock substrates. Its current distribution, cornered in the NW of the Northern part of the Montes de León, suggests a possible competitive exclusion between this taxon and I. galani, as the galani haplotypes (NML-II) appear cornered in the most harsh and continental areas, speaking also about a, even in the past, very limited presence of this species in the area that probably was soon absorbed by I. m. astur ssp. nov. (with NML-Concluding, it seems that the current main distribution area of I. m. astur ssp. nov. (especially the typical NML-I) gravitates around what was the divisory between watersheds in the past, later shifted to the East during the Quaternary. Eastern known limits of I. m. astur ssp. nov. do not pass away from Collado de Campo Lamoso (1500 m), which today is perfectly suitable for the species, but during the Pliocene and the main part of the Pleistocene, constituted a barrier across which the two northern immediate valleys drained to the southern slopes. The West-East continuity of this massif during the end of the Miocene was broken by changes in the drainage across this pass in the Pliocene (geological datation uncertain). Although nowadays the pass to the East (to the Filera Massif, 1879 m) is possible for Iberolacerta, the prospections in these drier limestone areas had been unfruitful. In the north of these Sierras, the species can reach up to Cascaros peak (1854 m), but this extreme has to be confirmed.

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Two populations of Lacerta monticola cyreni from the sierra de Gredos (Avila, central Spain) have been studies. Eight egg-layings in captivity and one from natural habitat have been analyzed. Nineteen specimens finally hatched with represents a reproductive success of about th 40 %. Egg-laying occurs in July and the clutches range from 3 to 9 eggs, with an average of 5,4 eggs per clutch. Hatching begins around the last days of August and last until the end of September. The newborn females are larger in S.V.L. (snout-vent length) than males. The colour and pattern of the hatchlings is very similar within the same clutch, but these characteristics differ from one clutch to another. The degree of calcification of the eggshells along with the environmental humidity during the period of incubation have proved to be two important factors to take into consideration in order to obtein a high reproductive success.

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

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

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The low-altitude populations of the threatenedlacertid lizard Iberolacerta monticola in A Coruña province have suffered population declines in the last decades. The present study has been done on the main population of this species in the Mandeo river basin, located in the river reserve of Chelo. Population size and density have been estimated, as well as other population parameters. We used the MARK program, which allows to use different variants of the Jolly-Seber method. In addition, we did an analysis of the sex-ratio and the structure of the population using size classes.The results indicated a high density, but limited to a reduced area with favourable habitat, surrounded by inadequate zones (forests with dense tree coverage),as well asa high mortality of immature individualsand an unequal proportion betweenimmaturesand adults. It seems to point out that this population shows insular characteristics. Other relevant data forthe survival of this population were obtained, as the apparent lack of predators, based on the low percentage of individuals with tail loss, or the reduced presence of individuals affected by ectoparasites of the genus Ixodes. The low number of observed movements points out that the individuals of this population are essentially sedentary. Some consequences for the survival of this population are set and some measures are proposed for its conservation.

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Las nuevas técnicas de secuenciacón permiten la identificación de una gran cantidad de polimorfismos de un único nucleótido (SNPs), que pueden ser empleados para detectar el grado de variación en una muestra de individuos y aplicarse a estudios poblacionales. En este trabajo se analizaron 90 individuos del lacértido Iberolacerta monticola de la población presente en la ZEC Betanzos-Mandeo. Se empleó el pipeline de STACKS para obtener, a partir de las lecturas procesadas, un catálogo de loci, genotipar todos los individuos y llevar a cabo el análisis genético de la población. Previamente a la ejecución, se realizó una exploración del espacio paramétrico. Además, se elaboró una lista negra de loci para poder restringir los análisis poblacionales a loci autosómicos. El estudio concluye con la determinación de la estructura genética de la población, que permitió observar el grado de fragmentación y detectar barreras que pueden estar afectando a la supervivencia de la misma.

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

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

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Male reproductive and fat body cycles were studies both in a mountain and a lowland population of the lizard Lacerta monticola cantabrica. Specimens were collected at two localities in Asturias (Northern Spain): Vegadeo (100 m a.s.l.) and Aramo (1450 m a.s.l.). General patterns of spermatogenesis in both populations were similar, and agree with the mixed type proposed by Saint Girons (1963). Spermatogenesis and testicular growth start in August and spermiogenesis begins in early autumn; before hibernation (October) spermiogenesis ceases earlier in the mountain than in the lowland population. The culminative phase of spermatogenesis and the testis weight peak occur after the onset of spring activity; there is then a progressive decrease that lasts until the end of the female`s ovulation cycle in July. Recrudescence of epididymides occurs mainly in late winter-early spring, with a pronounced development just after emergence from wintering; there is a delay in the epididymis peak of the mountain population. Both populations show an abrupt epididymal decline in the summer, between June and August. Fat bodies begin to increase in summer, just after the end of the reproductive period, reaching maximum weights in autumn prior to wintering. Pre-hibernation fat body weights for males at the two elevations are nearly the same (about 2% of body weight on average), but winter consumption is much larger in the lowland population in which there is some activity during this period. Fat bodies are not depleted at emergence and the remaining stores are utilized during the spring reproductive period, leading to minimum levels in June in both populations.

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Within ten years, data on twenty amphibian and thirty reptile species from the Iberian Peninsula were collected. The excursions were mainly centred on the northeast of Spain and the Extremadura. We report ob- servations on Salamandra salamandra almanzoris MÜLLER & HELLMICH, 1935, Rana pereti SEOANE, 1885, Lacerta viridis (LAURENTI, 1768), Lacerta monticola BOULENGER, 1905, and Malpolon monspes- sulanus (HERMANN, 1804). Additionally, all herpetological records and brief descriptions of the correspond- ing habitats are listed.

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A phylogenetic-comparative approach was used to assess and refine existing secondary structure models for a frequently studied region of the mitochondrial encoded large subunit (16S) rRNA in two large lizard lineages within the Scincomorpha, namely the Scincidae and the Lacertidae. Potential pairings and mutual information were analyzed to identify site interactions present within each lineage and provide consensus secondary structures. Many of the interactions proposed by previous models were supported, but several refinements were possible. The consensus structures allowed a detailed analysis of rRNA sequence evolution. Phylogenetic trees were inferred from Bayesian analyses of all sites, and the topologies used for maximum likelihood estimation of sequence evolution parameters. Assigning gamma-distributed relative rate categories to all interacting sites that were homologous between lineages revealed substantial differences between helices. In both lineages, sites within helix G2 were mostly conserved, while those within helix E18 evolved rapidly. Clear evidence of substantial site-specific rate variation (covarion-like evolution) was also detected, although this was not strongly associated with specific helices. This study, in conjunction with comparable findings on different, higher-level taxa, supports the ubiquitous nature of site-specific rate variation in this gene and justifies the incorporation of covarion models in phylogenetic inference.

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The rock lizard Lacerta monticola is a climatic relict confined to three mountain ranges and one coastal region of the Iberian peninsula. Scalation was studied in specimens from localities encompassing all major parts of its range. Population differentiation was analysed using analyses of variance, z-transformations of within-locality character-correlations, canonical variate analyses, principal components analysis, and matrix association tests. Substantial differentiation was found between Pyrenees specimens and those from other localities, supporting a recent study which indicated that the Pyrenees populations could represent a different species. Considerable geographic variation is also found among populations from the remaining parts of its range, the pattern of which is not compatible with existing racial categories designated for this species.

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Sexual selection favors more conspicuous male displays, whereas natural selection (via predator pressure) favors less conspicuous displays. However, this trade-off might be altered if males with more conspicuous displays could compensate behaviorally for their increased conspicuousness by acting more cautiously toward predators. The aim of this study was to explore in 2 species of Iberian rock lizards whether or not conspicuous coloration was associated with antipredatory behavior and whether conspicuousness-dependent regulation of antipredatory behavior existed. Our results suggested that male lizards may compensate for the negative effects of conspicuous sexual coloration on predation risk by modulating their antipredatory behavior (time inside refuges, false alarms, etc). We found that male Iberolacerta monticola, but not male Iberolacerta cyreni, compensated for the negative effects of blue lateral ocelli, which increased visual conspicuousness. However, male lizards did not compensate for relatively unexposed ventral spots. We also found that male I. monticola in better condition and with more blue lateral ocelli were shier, whereas male I. cyreni in better condition and with more ventral spots were bolder. These 2 lizard species live in habitats that differ in refuge availability and in the number of potential predators, which may promote differences in the trade-off between predation risk and social behavior and may explain the observed interspecific differences in antipredatory behavior. This suggests that regulation of antipredatory behavior may also function as a condition-dependent cost promoting costly (honest) sexual signaling in some species.

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

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West European Rock lizards, Lacerta (Iberolacerta) have small widely separated ranges in highland areas. Mitochondrial and nuclear DNA sequences corroborate the monophyly of the group and show it is not closely related to any of the other Rock lizards with which it was formerly placed in Archaeolacerta, an assemblage for which there is no evidence of clade status. L. (Iberolacerta) consists of four main units: L. (I.) horvathi of NW Croatia and neighbouring regions; the Pyrenees species, L. (I.) bonnali, L. (I.) aranica and L. (I.) aurelioi; L. (I.) cyreni of the Iberian Sistema Central, with distinctive populations in the Sierras de Béjar,Gredos and Guadarrama; and L. (I.) monticola of the Serra da Estrela of Central Portugal and NWSpain, this unit also contains L. (I.) cyreni martinezricai of La Peña de Francia, W. Spain and a distinctive population in the Montañas de Sanabria. L. (Iberolacerta) has persisted in some mountain ranges for at least 4.2±1.4Ma and may have been restricted tomountains by competition from Wall lizards (Podarcis). Its clade status shows it has lost range extensively and has produced few external branches since its initial fragmentation. In contrast, Podarcis diversified about the time L. (Iberolacerta) fragmented, producing a series of widespread lineages that have persisted until the present time. The mainly European subfamily Lacertinae, to which both L. (Iberolacerta) and Podarcis belong, diversified rapidly 13–9 Ma ago, probably largely replacing other lacertid lizards of earlier origin. As another round of replacement started at approximately 9 Ma ago with the spread of Podarcis, this may be a recurrent phenomenon in the evolution of some lizard communities.

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This paper presents an observational and experimental study of the basking behaviour and heat exchange rate of the montane lizard Lscerta monticola. The results obtained by these procedures were coupled in order to understand behavioural mechanims promoting effective thermoregulation at high altitudes. Heating rate was higher when body size was smaller, and substrate temperature and sun rays incidence angle were higher. The lizards cooled faster when body size and substrate temperature were lower, and when the body temperature of the lizard going into shadow was higher. Time exposed to sun and mean duration of basking periods were longer early in the morning, while bask frequency increased through the morning. Our results suggest that time devoted to basking is mainly obtained by regulating bask duration. Lizards obtained the necessary time for heating by means of long basking periods. Mean travel distance per minute and distance to the nearest refuge increased from early morning to midday. These behavioural variables were tightly correlated with the expected heating rate of individuals. Body size affects thermoregulatory behaviour as well as locomotor activity. Juvenile lizards, with small body mass and high surface-to-volume ratios, were subjected to faster heating and cooling rates, basked more frequently than adults (but during shorter periods), and devoted more time to locomotion than adults. The thermoregulatory behaviour of L. monticola is the result of the combination of shuttling heliothermy by basking and the exploitation of thermal opportunities offered by patches in shade through thermal exchange with the substrate.

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

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We discuss three aspects of the thermal biology of lacertid lizards. First, we provide an overview of the available data on field body temperatures (Tb), the thermal sensitivity of various performance functions and selected body temperatures in different species of lacertid lizards. We also briefly summarise information on the mechanisms of thermoregulation. Second, we discuss recent developments to estimate the »precision« of thermoregulation, and the contribution of distinct behavioural mechanisms. Finally, we revise available evidence for the existence of evolutionary adjustments of thermal characteristics in lacertid lizards. Existing studies have mainly dealt with within- and among-species differences in thermoregulatory behaviour (selected temperatures) and thermal physiology of adults (optimal temperatures, heating rates). Available data provide only limited evidence for clear-cut evolutionary shifts in thermal physiology characteristics along climatic gradients.

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Uncertainty in projections of global change impacts on biodiversity over the 21st century is high. Improved predictive accuracy is needed, highlighting the importance of using different types of models when predicting species range shifts. However, this is still rarely done. Our approach integrates the outputs of a spatially-explicit physiologically inspired model of extinction and correlative species distribution models to assess climate-change induced range shifts of three European reptile species (Lacerta lepida, Iberolacerta monticola, and Hemidactylus turcicus) in the coming decades. We integrated the two types of models by mapping and quantifying agreement and disagreement between their projections. We analyzed the relationships between climate change and projected range shifts. Agreement between model projections varied greatly between species and depended on whether or not they consider dispersal ability. Under our approach, the reliability of predictions is greatest where the predictions of these different types of models converge, and in this way uncertainty is reduced; sites where this convergence occurs are characterized by both current high temperatures and significant future temperature increase, suggesting they may become hotspots of local extinctions. Moreover, this approach can be readily implemented with other types of models.

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Amoeba associated microorganisms (ARMs) are bacteria or viruses that share a symbiotic relationship with amoebas. Many ARMs are associated with human diseases and it has been reported the acquisition of resistance inside their host. These facts highlight the importance of finding and characterizing ARMs in a public health’s perspective. In the present work, amoebas from environmental samples have been isolated and identified by optical microscopy and molecular methods. Amoeba associated viruses, especially from the NCLDV group, were also subjected to a PCR screening. From the 12 collected samples, it was possible to isolate 17 amoebas, of which 11 were identified by molecular methods. No viruses were detected but in 2 of these amoebas, a bacterium was identified which shares an 89% sequence identity with the Marivirga tractuosa species. If amoebas allow the replication of a Mimivirus, which belongs to the NCLDV group, would they be able to replicate iridoviruses that belong to the same group? An Acanthamoeba castellanii cell line was experimentally inoculated with three Ranavirus: FV3 (Frog Virus 3), Ma3B and BoA (respectively, a Triturus marmoratus and a Triturus boscai isolate). The PCR test suggested an association between A. castellanii cells with the Ma3B and BoA viruses. VLPs (Virus-like particles) were visualized by electron microscopy in the Ma3B inoculated amoebas. Ranavirus from several Triturus species are currently being characterized and the sequences of their Major Capsid Protein and DNA polymerase genes are identical. Could the sequencing of a less conserved gene like the Flap endonuclease, allow the distinction? The sequencing and analysis of the Flap gene was able to group viruses FV3 and LMO (Lacerta monticula virus) as they present an equal gene sequence. The three Triturus marmoratus isolates presented an equal Flap gene amongst them and were also grouped. Between the two groups, this gene sequence has a difference of 8 nucleotides, confirming the separation of the FV3 and LMO viruses from the other iridoviruses tested.

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Research on fluctuating asymmetry (FA)-mediated sexual selection has focused almost exclusively on visual signals and ignored chemical communication despite the fact that many species rely on chemical signals for attracting mates. Female mate choice based on visual traits appears to be rare in lizards. However, the femoral glands of male lizards produce pheromones which might transmit chemical information about an individual`s developmental stability. Therefore, we hypothesized that mate choice may be based on chemical cues. We analysed the effect of the developmental stability levels of males on the attractiveness of males` scents to females in a laboratory experiment with the lizard Lacerta monticola. When we offered two males of similar body size, females preferentially associated with the scents of males with low FA in their femoral pores and also with the scents of males with a higher number of femoral pores. This suggested that the females were able to discriminate the FA of the males by chemical signals alone and that the females preferred to be in areas marked by males of high quality, thus increasing their opportunities of mating with males of high quality. We suggest that the quality and/or amount of male pheromones could communicate the heritable genetic quality of a male to the female and thereby serve as the basis for adaptive female choice in lizards.

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

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

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

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The Iberian rock lizard (Iberolacerta monticola) has gone extinct or severely declined in several low-altitude and isolated populations in Galicia during the last three decades. In order to assess the conservation status of this species in NW Iberia, we recorded morpho- and eco- logical data of individuals from eleven localities of this area. In addition, the genetic variability of these specimens was screened by combi- ning nuclear (microsatellite) and mitochondrial markers (470 bp from the control region). The population established at O Pindo (A Coruña), at least 100 km apart from other populations, shows a conspicuous diagnostic difference in its mitochondrial lineage and the largest diffe- rentiation at microsatellite loci, as well as some morphological differentiation most likely linked to the particular habitat of this site, a relatively arid and rocky environment.

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

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

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

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DNA sequences from parts of the 12S, 16S and cytochrome b mitochondrial genes, which totalled 1049 aligned base pairs, were used to estimate the relationships of 49 species of Lacertidae, including representatives of 19 out of the 23 recognized genera and 23 species of the paraphyletic genus Lacerta. These data were used, together with morphological information, to estimate the relationships within the family. Molecular evidence corroborates the monophyletic status of many genera and species groups originally based on morphology. It indicates that Psammodromus forms a clade with Gallotia, which is the sister taxon of all other lacertids. These comprise three units: the primarily Afrotropical armatured group; the largely Oriental Takydromus; and the west Palaearctic Lacerta and its derivatives, Podarcis and Algyroides. Morphology also supports the first three assemblages, but suggests that they are derived from a paraphyletic Lacerta. Within Lacerta and its allies, DNA sequence analysis corroborates the affinity of some members of each of the subgenera Lacerta s. str. and Timon, and of the L. saxicola group. It also supports the relationship of L. monticola, L. bonnali and L. horvathi, and suggests that the L. parva–L. fraasi clade and L. brandti are not related to Psammodromus–Gallotia, as morphology indicates, but instead are associated respectively with the L. danfordi and L. saxicola groups. DNA sequence data provide additional evidence that the eastern Arabian `Lacerta` jayakari and `L.` cyanura are members of the armatured clade and also sister species. Our analysis supports an origin for present lacertids in west Eurasia. The armatured clade invaded Africa, probably in the mid-Miocene, spreading widely and evolving increasingly xeric-adapted forms, one lineage of which later moved back into the Palaearctic. `Lacerta` jayakari and `L.` cyanura are assigned to Omanosaura, Lutz and Mayer 1986. The name Gallotiinae Cano, Baez, Lopez-Jurado and Ortega, 1984 is available for the Gallotia–Psammodromus clade, Eremiainae Shcherbak 1975 for the armatured clade and Lacertinae for Lacerta, Podarcis and Algyroides. Two new subgenera of Lacerta are proposed here: Caucasilacerta for L. saxicola and its allies, and Parvilacerta for L. parva and L. fraasi.

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

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The evolutionary diversification of many terrestrial vertebrate groups is strongly linked to climatic events in the Cenozoic, the period from 65 Million years ago to today when modern animals first appeared. I investigated the effects of Cenozoic climate change on the taxonomic and morphological diversification of the Old World lizard family Lacertidae, with particular emphasis on the African radiation. African lacertids exhibit an unusual pattern of diversification, in which their highest species richness occurs in deserts north and south of the equator, despite being spread throughout the continent. This disparity is particularly surprising given that desert lacertids are thought to be evolutionarily younger than their mesic-dwelling relatives, suggesting increased diversification rates in arid habitats. To identify the evolutionary factors underlying this pattern, I use a combination of phylogenetic, morphological and ecological techniques. In Chapter 1, I apply Bayesian methods and fossil-based calibrations to molecular sequence data to construct a time-calibrated phylogeny for Lacertidae. I estimate that the family arose in the early Cenozoic, with the majority of their African radiation occurring in the Eocene and Oligocene. In Chapter 2, I describe changes in lacertid body shape across biomes and substrates, and find widespread morphological convergence in similar habitat types. I suggest that in addition to foraging demands, fluctuating and extreme climatic conditions, largely driven by precipitation and temperature, contribute to morphological convergence across independent arid-dwelling clades. Finally, I test if ancestral transitions in ecology, morphology, and rates of diversification temporally coincide with paleoclimatic events in the Cenozoic. I use High Resolution X-ray Computed Tomography to characterize changes in the skull related to life in arid habitats, and apply maximum likelihood methods to test if the origins of those traits temporally coincide with significant shifts in habitat, diversification rates and climatic changes. My results show that African lacertids experienced three major peaks in diversification, accompanied by the evolution of suites of arid-adapted morphological traits. These changes coincide with climatic shifts in Africa, including the transition from closed forests to open grasslands and savanna in the late Oligocene, prior to the peak temperatures of the mid-Miocene Climatic Optimum, and following the formation of the Benguela current leading to hyper-aridity in southern Africa. I conclude that deserts are important centers for reptile evolution, but that expected changes in climate due to global warming may outpace the ability of arid-dwelling species to adapt and persist in the future.

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Background: Although current molecular clock methods offer greater flexibility in modelling evolutionary events, calibration of the clock with dates from the fossil record is still problematic for many groups. Here we implement several new approaches in molecular dating to estimate the evolutionary ages of Lacertidae, an Old World family of lizards with a poor fossil record and uncertain phylogeny. Four different models of rate variation are tested in a new program for Bayesian phylogenetic analysis called TreeTime, based on a combination of mitochondrial and nuclear gene sequences. We incorporate paleontological uncertainty into divergence estimates by expressing multiple calibration dates as a range of probabilistic distributions. We also test the reliability of our proposed calibrations by exploring effects of individual priors on posterior estimates. Results: According to the most reliable model, as indicated by Bayes factor comparison, modern lacertids arose shortly after the K/T transition and entered Africa about 45 million years ago, with the majority of their African radiation occurring in the Eocene and Oligocene. Our findings indicate much earlier origins for these clades than previously reported, and we discuss our results in light of paleogeographic trends during the Cenozoic. Conclusion: This study represents the first attempt to estimate evolutionary ages of a specific group of reptiles exhibiting uncertain phylogenetic relationships, molecular rate variation and a poor fossil record. Our results emphasize the sensitivity of molecular divergence dates to fossil calibrations, and support the use of combined molecular data sets and multiple, well-spaced dates from the fossil record as minimum node constraints. The bioinformatics program used here, TreeTime, is publicly available, and we recommend its use for molecular dating of taxa faced with similar challenges.

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Environmental conditions experienced by a species during its evolutionary history may shape the signals it uses for communication. Consequently, rapid environmental changes may lead to less effective signals, which interfere with communication between individuals, altering life history traits such as predator detection and mate searching. Increased temperature can reduce the efficacy of scent marks released by male lizards, but the extent to which this negative effect is related to specific biological traits and evolutionary histories across species and populations have not been explored. We experimentally tested how increased temperature affects the efficacy of chemical signals of high- and low- altitude populations of three lizard species that differ in their ecological requirements and altitudinal distributions. We tested the behavioral chemosensory responses of males from each species and population to male scent marks that had been incubated at one of two temperatures (cold 16°C or hot 20°C). In high- altitude populations of a mountain species (Iberolacerta monticola), the efficacy of chemical signals (i.e., latency time and number of tongue flicks) was lower after scent marks had been exposed to a hot temperature. The temperature that scent marks were incubated at did not affect the efficacy of chemical signals in a ubiquitous species (Podarcis muralis) or another mountain species (I. bonalli). Our results suggest that specific ecological traits arising through local adaptation to restricted distributions may be important in determining species vulnerability to climatic change

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Immunological reactions tested according to Libby`s photronreflectometric technique show that: (i) Archaeolacerta Mertens, 1921, and Podarcis Wagler, 1830, are independent genera and not subgenera of Lacerta Linnaeus, 1758; (ii) Podarcis muralis (Laurenti, 1768), Podarcis sicula (Rafinesque, 1810), Archaeolacerta bedriagae (Camerano, 1885), Archaeolacerta monticola (Boulenger, 1905), Lacerta lepida Daudin, 1802, Lacerta schreiberi Bedriaga, 1878 and Lacerta viridis (Laurenti, 1768) are full species; (iii) Podarcis tiliguerta is a subspecies of P. muralis: P. muralis tiliguerta (Gmelin, 1789).

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Trade-offs between reproduction and survival are important determinants of life-history characteristics of lizards. Organisms cannot increase the allocation of limited resources to reproduction without diverting a proportional amount of energy from another trait. Locomotor performance is an ecologically relevant trait that potentially influences survival by affecting the ability to escape from predators. Most studies have used female lizards as subjects because pregnancy is known to reduce their locomotor abilities, whereas little is known on costs of reproduction in males. In this study we suggest that in males of the lizard Lacerta monticola reproductive investment in morphological traits that confer dominance (i.e. head size) might lead to a low probability of survival by decreasing investment in other traits that affect locomotor performance (i.e. limb symmetry). We staged laboratory agonistic encounters between males and measured their morphology and burst speed on a race track to examine possible relationships between morphology, social dominance and locomotor capacity. Our results indicate that social dominance was positively related to relative head height, and that escape speed was negatively related to levels of fluctuating asymmetry in femur length, but also negatively related to relative head height. Males with greater relative head height also had more asymmetrical femurs, thus dominant males suffered a decrease in locomotor performance. Males with higher heads tend to dominate male–male interactions and hence may gain access to reproductive females, thus increasing their current reproduction success. However, this might occur at the expense of future survivorship mediated by a decrease in escape speed. Therefore, in male L. monticola there might be a trade-off between current reproductive success and survival.

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In spite of the importance of chemoreception in social organization and sexual selection of lizards, there is a lack of general knowledge on how the characteristics of chemical signals mediate these behaviors. Moreover, it is unknown which are the mechanisms that might confer honesty to the information provided by chemical signals. We analyzed here whether characteristics of the lipophilic fraction of femoral gland secretions of Lacerta monticola male lizards can be related to the morphology, physical condition, and health state of the sender. Our results indicated that some male traits, such as body size, number of blue spots, and number of femoral pores and their level of fluctuating asymmetry, were related to variability in the relative proportions of some lipophilic chemical compounds found in secretions. Thus, conspecifics could obtain reliable information on the producer of a scent mark based on chemicals alone, and this might be the basis of female choice observed in this lizard. Moreover, only males with a greater T-cell immune response had higher proportions of two steroids (ergosterol and dehydrocholesterol) in their femoral secretions, which might suggest that the signal is honest and costly to produce. We suggest that only high-quality males could divert these compounds from metabolism to secretions in order to produce an exaggerated and honest `chemical ornament.`

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Female mate choice based on visual traits appears to be rare in lizards. Field observations suggest that females of the lizard Lacerta monticola preferred to mate with larger/older males. Although older males are usually green and larger, and younger males brown and smaller, there is some overlap in size and coloration between age classes. Thus, visual cues may not always be reliable indicators of a male`s age. We hypothesized that female mate-choice preferences may be based on males` pheromones, which might transmit information about characteristics such as age. In a laboratory experiment, we analyzed the effect of age of males on attractiveness of their scents to females. When we offered scents of two males of different age, females associated preferentially with scents of older males. This suggested that females were able to assess the age of males by chemical signals alone, and that females preferred to be in areas scent-marked by older males. Thus, females may increase their opportunities to mate with males of high quality, or may avoid harassment by sneaking young males. This result agreed with field observations on females mating with old males, and rejection of advances by young males. Our results also suggested that female preference for older males may depend on their own body size. Large females showed a strong preference for older males, whereas smaller females were not so selective. This, together with males` preference for large females, might lead to size-assortative matings. We suggest that the quality and/or quantity of male pheromones could communicate to the female heritable male genetic quality (i.e. age) and thereby serve as the basis of adaptive female choice in lizards.

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Individual animals from the same population, sex, age and reproductive status often respond differently and consistently to predators. One important dimension of this behavioural variation is the shy–bold continuum. Innate differences in boldness might explain why individuals differ in their antipredator behaviour. In a laboratory experiment, we examined the sources of individual variation in antipredator behaviour of adult male lizards Lacerta monticola. We simulated in the laboratory repeated predatory attacks of low or high risk and analysed activity levels and refuge use in both situations. Multivariate analyses suggested the existence of two consistent and independent shy–bold continua. The first described a gradient from bold lizards that spent shorter times in the refuge after predatory approaches to shy lizards with longer emergence times, whereas the other described a gradient from bold lizards with a low propensity to hide when the predator was close but risk was low to shy lizards that hid more often. We analysed whether morphological characteristics, body condition and health (estimated from their T cell immunocompetence) of individuals might account for the differences observed. Bold individuals had smaller absolute body size, but relatively larger heads, better body condition and better health. Bold individuals with a low propensity to hide when risk was low had larger absolute body sizes, whereas relative head size, and body condition and health were not important. We suggest that the position of an individual in the shy–bold continua might reflect its optimal antipredator behaviour, which would be a function of its health, general quality and ability to evade predators.

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Vertebrate males often show breeding colours that may function as reliable signals of status in intrasexual competition. In many lacertid lizards, males show a conspicuous row of small but distinctive blue spots that runs along their body side on the outer margin of the belly. However, no study has examined the role of these blue spots. We first analysed in a field population of the Iberian rock lizard, Lacerta monticola, the relationships between number of blue spots and some morphological traits, which are known to be related to males’ fighting ability. The number of spots seems to be an character showing ontogenetic change as large (generally older) males showed more blue spots than small (generally younger) males. Males with a higher body condition also showed a higher number of blue spots. Thus, a higher number of blue spots may be used to signal size, age or body condition. Many contiguous blue spots would result in a visual artefact consisting of a continuous blue band, which might be a reliable size- or condition-dependent signal in some social contexts. We further examined in the laboratory whether male characteristics are related to dominance status. In males with similar body size or age, those with relatively larger heads were more dominant, whereas the number of blue spots was not important. Moreover, the number of blue spots in nature was not related to relative head size. Finally, we experimentally manipulated the presence and the number of blue spots of intruding males, and examined the aggressive response of resident males. Intruder individuals manipulated to cover all their blue spots received a lower amount of aggression. However, males with different numbers of manipulated blue spots received a similar number of aggressive responses. These results suggest that, during agonistic encounters, the presence of blue spots, but not their number, may elicit aggressiveness. Thus, blue spots may serve to identify an individual as an adult male, and to enhance body size of larger males.

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In the lizard Iberolacerta monticola, there are 2 discrete color morphs (`blue` vs. `green`) that seem to have alternative reproductive strategies. Because scent marking of territories and chemoreception are important in social organization of these lizards, we explored whether there is also chemical polymorphism and chemosensory recognition between color morphs. Analyses by gas chromatography-mass spectrometry showed that adult males of different morphs had similar chemical compounds in femoral gland secretions, but the proportions of some shared chemicals were different; blue morph males had higher proportions of steroids and lower proportions of fatty acids than green males. Differential tongue-flick rates to scents from femoral secretions of males indicated that males were able to detect the scent of any male from a baseline odorless control (water) but did not discriminate between the scent of males of different morphs. In contrast, females detected and discriminated between scents of the 2 male morphs by chemical cues alone. We suggest that differences in males` chemical signals between morphs and the ability of females of discriminating chemically between male morphs might be required to maintain spatial behavior, social organization, and mate choice decisions and, thus, be important for the maintenance of alternative reproductive strategies and polymorphism in lizards.

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Female preference for dominant males is widespread and it is generally assumed that success in male-male competition reflects high quality. However, male dominance is not always attractive to females. Alternatively, relatively symmetric individuals may experience fitness advantages, but it remains to be determined whether males with more symmetrical secondary sexual traits experience advantages in both intra- and intersexual selection. We analysed the factors that determine dominance status in males of the lizard Lacerta monticola, and their relationship to female mate preference, estimated by the attractiveness of males` scents to females. Sexually dimorphic traits of this lizard (head size and femoral pores) appear to be advanced by different selection pressures. Males with relatively higher heads, which give them advantage in intrasexual contests, were more dominant. However, head size was unimportant to females, which preferred to be in areas marked by relatively heavier males, but also by males more symmetric in their counts of left and right femoral pores. Chemicals arising from the femoral pores and other glands might honestly indicate quality (i.e. related to the symmetry levels) of a male to females and may result from intersexual selection. Females may use this information because the only benefit of mate choice to female lizards may be genetic quality. Chemical signals may be more reliable and have a greater importance in sexual selection processes of lizards than has previously been considered.

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ln der vorliegenden Arbeit wird die Herpetofauna von über 50 portugiesischen Gebirgszügen zusammengestellt: für jedes Gebirge der hõchste bisher bekannte Fundort der einzelnen Arten mit Hôhenangabe. Es ist das Ergebnis vierjiähriger Untersuchungen (1976-1980).

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

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Prey often respond to predators by increasing their use of refuges but relatively few studies have analyzed how prey decide when to resume their behavior after a predator’s unsuccessful attack. This is important because refuge use may have some costs that should be minimized such as the loss of time available for foraging or mate searching. In addition, unfavorable conditions in refuges (e.g., suboptimal temperatures) might entail physiological costs such as hypothermia. Under these circumstances, animals should optimize the decision of when to come out from a refuge by balancing the fitness effects of the diminution of predation risk with time against the costs of loss of time available for other activities and loss of time spent at optimal body temperature. I review several experiments with lizards that support that individuals decide to come out from a refuge when the costs of hiding exceed predation risk in the exterior, and that there is an optimal emergence time. Optimization of these antipredatory behavioral strategies might help lizards to cope with changes in predation risk without incurring excessive costs.

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Prey often respond to predator presence by increasing their use of refuges. However, unfavorable thermal conditions in refuges might entail physiological costs for an ectothermic prey. Thus, the decision of when to come out from a refuge should be optimized by considering the expected fitness effects of diminution of predation risk with time, but also by considering the cost of the loss of time spent at optimal body temperature maximizing physiological functions. The model of Ydenberg and Dill describes the trade-off between risk and cost for a prey fleeing to a refuge. We present a special case of this model to predict how emergence time from the refuge in lizards or other ectotherms should vary as a function of risk of predation and thermal costs of refuge use. The analyses of the variation in emergence time from a refuge of Lacerta monticola lizards in the field under two different predation risk levels supported the predictions of the model. As predicted, time spent in the refuge was longer when the threat of the initial attack had been higher, and therefore the subsequent diminution of risk was slower, but only when lizards emerged at the same place where they hid. When initial body temperature was high, some lizards decreased emergence time by emerging from a different place. In addition, the effects of thermal costs were more relevant in the high-risk situation. Time spent in the refuge under high risk increased when thermal conditions of the refuge were more similar to thermal conditions outside (i.e., physiological costs of refuge use were lower). We conclude that optimization of refuge-use strategies might help lizards cope with changes in predation risk without incurring excessive physiological costs.

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Theoretical models of anti-predator escape behaviour suggest that prey may adjust their escape response such that the optimal flight distance is the point at which the costs of staying exceed the costs of fleeing. Anti-predatory decisions should be made based also on consequences for long-term expected fitness, such as the costs of refuge use. For example, in lizards, the maintenance of an optimal body temperature is essential to maximize physiological processes. However, if unfavourable thermal conditions of refuges can decrease the body temperature of lizards, their escape decision should be influenced by refuge conditions. Analyses of the variation in flight distances and emergence latency from a refuge for the lizard Lacerta monticola under two different predation risk levels, and their relationship with the thermal environment, supported these predictions. When risk increased, lizards had longer emergence latencies, and thus costs of refuge use increased (a greater loss of time and body temperature). In the low-risk situation, lizards that were farther from the refuge had longer flight distances, whereas thermal conditions were less important. When risk increased, lizards had longer flight distances when refuges were farther off, but also when the external heating rate and the refuge cooling rate were lower. The results suggest that, in addition to the risk of predation, expected long-term fitness costs of refuges can also affect escape decisions.

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Not all individual lizards in a population are simultaneously active even when thermal conditions are suitable for activity. We examined inter-individual variability in activity levels of male Iberian rock lizards (Lacerta monticola) in a seminatural enclosure during the mating season, and analyzed whether social status affects their activity levels, time budgets, and body-mass changes. Activity levels of lizards varied significantly with time of day. However, activity levels of individual males were significantly correlated with their rank in the social hierarchy. When the males were active, their status did not influence the time spent basking, resting, or moving, but males with a higher status spent more time in social activities. Higher activity levels were costly, causing males to lose more body mass, although this could have been mainly due to the costs of maintaining a higher social status. We conclude that because attaining a higher status may require a male to be more active and more involved in agonistic encounters, subordinate individuals decrease their activity in order to decrease the costs of social behavior.

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Prey often respond to predator presence by increasing their use of refuges. However, because the use of refuges may entail several costs, the decision of when to come out from a refuge should be optimized. In some circumstances, if predators remain waiting outside the refuge and try new attacks or if predator density increases, the prey may suffer successive repeated attacks in a short time. Successive attacks may represent an increase in the risk of predation, but the costs of refuge use also may increase with time spent in the refuge. Thus, prey should make multiple related decisions on when to emerge from the refuge after each new attack. We simulated in the field repeated predatory attacks to the same individuals of the lizard Lacerta monticola and specifically examined the variation in successive times to emergence from a refuge under different thermal conditions (i.e., different costs of refuge use). The results showed that risk of predation but also thermal costs of refuge use affected the emergence decisions. Lizards increased progressively the duration of time spent in the refuge between successive emergence times when the costs of refuge use were lower, but tended to maintain or to decrease the duration of time spent in the refuge between successive emergence times when cost of refuge use increased. Additionally, lizards that entered the refuge with higher body temperatures had overall emergence times of longer duration. Optimization of refuge use and flexibility in the antipredator responses might help lizards to cope with increased predation risk without incurring excessive costs of refuge use.

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In lizards, ontogenetic changes in body size affect thermal-exchange rates. This simple physical property may have consequences for thermoregulation, and also for antipredator behavior. We examined how ontogenetic changes in body mass affect rates of heating and cooling of the lizard Lacerta monticola, confirming the general result obtained for other lizards. We further analyzed the differences between juveniles and adults in approach distances to a simulated predator and in time to emerge from refuges. Juvenile lizards have a lower absolute running speed, making them more vulnerable to predation. However, in contrast to results expected from optimal-escape theory, approach distances were shorter for juveniles than for adults. Juveniles may be confident in their small size and only flee when the probability of being detected is high. On the other hand, differences in thermal properties might affect costs of refuge use. Thus, juveniles might delay fleeing because their costs of hiding are higher, as they cool faster than adults. Differences in thermal costs may also explain the juveniles` shorter times of emergence from refuges. Because of the behavioral adjustments involved in antipredator behavior, the physiological costs of reaching a low body temperature in refuges probably do not differ between age classes.

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Prey often respond to predator presence by increasing their use of refuges, but because this strategy may be costly, the decision regarding when to come out from a refuge should be optimized. The loss of foraging opportunities may be one of the main costs when safer microhabitats (i.e. refuges) are also the poorest in terms of their foraging profitability. We present the results of an experimental field study to test whether emergence times from a refuge of the Iberian rock lizard, Lacerta monticola, vary as a function of expected foraging opportunities and level of satiation of the lizard. As predicted, short-term fluctuations in availability of food influenced emergence times; when a lizard had just detected some food in the recent past, emergence times decreased greatly, because the loss of opportunities for foraging increased costs of refuge use. Furthermore, the characteristics and success of the encounter with food, nutritional state of lizards, and the added possibility of capturing new food items influenced the duration of hiding times. Therefore, foraging requirements and avoidance of predators may be conflicting demands that L. monticola lizards balance by modifying the duration of time spent in refuges.

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1 In spite of the fact that chemoreception plays an important part in sexual selection of many vertebrates, there is a lack of general knowledge on the role of specific chemical compounds as sexual signals, and on how the characteristics of chemical signals mediate behaviours such as mate choice. 2 We examined how female Iberian Rock Lizards use the chemical characteristics of femoral gland secretions of males to select potential mates. 3 Tongue-flick assays and choice of scent trials showed that females discriminated by chemosensory cues alone differences in chemical secretions of different males. Females showed stronger chemosensory responses and preferred the scent of males of presumably high quality (i.e. those more symmetric and with a greater immune response). 4 Chemical analyses by gas chromatography-mass spectrometry showed that these preferred males allocated relatively more cholesta-5,7-dien-3-ol and ergosterol to femoral secretions, which suggested that females used these chemicals to choose between males’ scents. 5 Further experiments confirmed that females were able to discriminate cholesta-5,7-dien-3-ol, ergosterol, and changes in their concentrations, from similar steroids (i.e. cholesterol) also found in secretions of males. Moreover, females were more attracted to areas that were experimentally manipulated to increase the proportion of ergosterol in natural scent marks of males. 6 We suggest that femoral secretions with higher proportions of cholesta-5,7-dien-3-ol and ergosterol might be reliable advertisements of quality of a male, which females could use to select mates.

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Evolutionary theory proposes that signals used in sexual selection can only be stable if they are honest and condition dependent. However, despite the fact that chemical signals are used by many animals, empirical research has mainly focused on visual and acoustic signals. Vitamin D is an essential nutrient for lizards, but in some lizards its precursor (cholesta-5,7-dien-3-ol=provitamin D) is found in femoral gland secretions, which males use for scent marking and intraspecific communication. By allocating provitamin D to secretions, males might need to divert vitamin D from metabolism. This might be costly and condition dependent. We tested whether diet quality affected chemical signals of male Iberian rock lizards (Lacerta monticola) and its consequences for sexual selection. After experimental supplementation of dietary vitamin D, males increased the proportion of provitamin D in femoral secretions. Further experiments showed that females detected these changes in males` signals by chemosensory cues, and discriminated provitamin D, and changes in its concentration, from similar steroids (i.e. cholesterol) found in secretions. Moreover, females preferred areas scent marked by males with more provitamin D in their secretions. This mechanism would confer honesty to chemical signals of male lizards, and, thus, females may rely on it to select high-quality males. We suggest that the allocation of vitamins and other essential nutrients to either visual (e.g. carotenoids) or chemical ornaments might be the common basis of honest sexual displays in many animals.

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Intrasexual competition favours the evolution of conspicuous fighting ability badges. However, in spite of the fact that chemoreception is important in sexual selection of many animals, such as lizards, the role of chemical signals in males` contests is relatively unknown. Here, we show that proportions of cholesterol in femoral gland secretions of male Iberian rock lizards were related to their body size (which confers a competitive advantage in fights). Males discriminated chemically and responded aggressively to cholesterol stimuli presented on swabs. Moreover, we experimentally increased cholesterol in the scent of males, and staged encounters in neutral cages between two unfamiliar and size-matched males. Focal males lost more agonisitic interactions against males manipulated with cholesterol than in control tests. We suggest that differences in scent composition may reliably signal fighting ability in many lizard species, which would help to avoid the costs of fighting.

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Because time spent in refuge may be costly if prey lose opportunities to forage, fight, or mate, prey allow predators to approach closer before beginning to flee when opportunity costs are high. Because the same opportunity costs may apply to refuge use as to escape, prey should make similar trade-offs between risk of emerging and cost of remaining in refuge. In the Iberian rock lizard, Lacerta monticola, we studied the effects of sex, reproductive season, speed of predator approach, and potential loss of mating opportunities on time spent in refuge following simulated predatory attacks. Lizards of both sexes adjusted refuge use to the level of risk by spending more time in refuge when approached rapidly than slowly. Females remained in refuge for equal times in the mating and postreproductive seasons, but males emerged sooner during the mating season, suggesting adjustment to a cost of lost opportunity to search for mates during the mating season. When a tethered female was nearby, males emerged from refuge earlier than if no female was present, indicating a trade-off between risk and mating opportunity. Approach speed affected emergence time when females were absent, but not when a female was present. Approach speed did not affect the probability that, after emerging, a male would return to court the female. For males that courted females intensely (bit them) before entering refuge, approach speed did not affect latency to emerge, but males that courted less intensely emerged sooner if approached slowly than rapidly. These findings show that males adjust the length of time spent in refuge to both risk of predation and reproductive cost of refuge use.

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We studied the relationships between thermoregulatory basking postures and heating rates in the Iberian rock lizard (Lacerta monticola). Heating rates were high when body size was small, the substrate temperature was high, and the angle of incidence of the sun`s rays was high (i.e., the rays were perpendicular to the lizard`s body). However, heating rates were not related to the compass orientation of lizard`s body axis to the sun (perpendicular versus in line with the sun). In the field, the slopes of the body axis of basking individuals were higher during the early morning. However, neither the cardinal orientation of a lizard`s body axis to the sun nor the frequency of flattening varied during the morning. Postural adjustments to select an adequate angle of incidence of the sun`s rays on the dorsal part of the body suggest that this behaviour might enhance absorption of solar radiation.

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1. Male competition for females often results in the evolution of conspicuous male traits that signal fighting ability or dominance status. Most studies examining mechanisms allowing these traits to function as reliable status signals (or badges) have focused on conspicuous visual or acoustic traits. However, many animals communicate chemically, and chemical traits alone may also signal male dominance status. 2. We examined whether chemicals in femoral gland secretions of male Iberian rock lizards ( Lacerta monticola monticola Boulenger 1905) may signal dominance status. 3. In staged encounters, larger males were dominant over smaller ones. While controlling for male body size, males of higher dominance status produced femoral secretions with higher proportions of hexadecanol and octadecanol. 4. Tongue-flick assays showed that males were able to (1) discriminate, by chemosensory cues alone, the different concentrations of hexadecanol from other chemicals presented in femoral secretions; (2) respond aggressively towards hexadecanol while responding neutrally towards other chemicals; and (3) show differential chemosensory and aggressive behaviours towards hexadecanol according to their own dominance status. 5. These results suggest that hexadecanol may be a reliable status badge. Moreover, because hexadecanol elicits male aggressive behaviour, subordinate males signalling high status (‘cheaters’) may end up paying high fighting costs. In addition, males that had higher dominance status, and that allocated higher proportions of hexadecanol to femoral secretions, had greater T-cell immune responses. This result suggests a possible link between quality of the immune system, dominance status, and chemical signals.

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Many lizards autotomize their tail to escape when attacked by a predator. The foraging tactics of tailless lizards may be affected by changes in behavior aimed at compensating tail loss per se as well as associated costs of regeneration. We tested this hypothesis in a population of the Iberian rock-lizard (Lacerta monticola), during the reproductive season, by comparing the diet of tailed and tailless lizards with the estimated prey availability in their habitat. We also compared the feeding rates, foraging microhabitat and foraging tactics of tailed and tailless lizards. Tail loss did not affect feeding rates, nor did it alter the foraging tactics of either sex during spring. Tailless lizards used rocky microhabitats where predation was avoided, but where food availability was lower, and thus diet diversity decreased at least among tailless males. Their diet consisted primarily of Bibio flies, an apparently suboptimal prey item.

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Many lizards autotomize their tails to escape when grasped by a predator. It is hypothesized that tail loss causes a reduction in social status, thereby potentially lowering their reproductive success. We experimentally induced tail loss in Lacerta monticola in a semi-natural enclosure, and show that tail loss reduced social status and mating access in males. Tailless males increased body mass more rapidly than tailed dominant males, probably due to lower aggression costs. Also, tailless females were courted less and copulated less than tailed females, supporting the hypothesis that tail loss decreases reproduction potential.

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The consequences of tail loss on the thermal biology of high-altitude rock lizards (Lacerta monticola) were studied by comparing field body temperatures, selected body temperatures in a gradient, and behavioural patterns of thermoregulation of tailed and tailless lizards in the field. Neither field nor selected body temperatures differed significantly between these groups. Time spent basking, mean duration of basking, and basking frequency did not differ between tailed and tailless lizards. Tailless lizards seemingly adjusted for running impairment by using rocks to a greater extent than did tailed lizards, and low morning body temperatures by increasing maximal duration of basking periods and keeping shorter distances to the nearest refuge. Tailless lizards modify thermoregulatory behaviour, but obtain the same thermoregulatory precision as tailed lizards, possibly allowing maximization of physiological processes linked to tail regeneration.

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Many lizards use caudal autotomy as an antipredatory strategy. We experimentally examined the effect of tail loss on the time-budgets, movement patterns, and home range size of the Iberian rock lizard Lacerta monticola in the field during the mating season. Our results indicate that tail loss did not alter most variables in male and female time-budgets but did affect the time spent moving and movement patterns of males. Tailless males spent less time moving and, when moving, more time on rocks, with shorter movements, more pauses, and less time on bushes than did tailed ones. Home range size was smaller in tailless males but did not differ between tailed and tailless females. The results suggest that tailless males may compensate the survival cost of autotomy by modifying their use of space and time; as a result, however, they may incur lower access to females.

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Parts of the mitochondrial genes coding for 12SrRNA and 16SrRNA (together about 960 bp) were sequenced for all Mediterranean species of `Mountain lizards` of the genera Archaeolacerta (sensu lato) and Iberolacerta. All subspecies of the Iberian species Iberolacerta cyreni and I. monticola were included in this study. In addition, samples of Apathya cappadocica and Darevskia rudis were analysed to elucidate the relationships of the European `Mountain lizards` to their possible relatives in the Near East. Maximum parsimony and neighbour joining analyses lead to the following major conclusions: (i) the monophyly of the genus Iberolacerta is very well supported; (ii) Archaeolacerta bedriagae (the type species of the genus) is most basal with respect to the ingroup taxa. If we accept Iberolacerta as a genus, Archaeolacerta becames paraphyletic. Therefore, we propose to restrict Archaeolacerta to the type species and to treat A. mosorensis and A. oxycephala provisionally as members of the collective genus Lacerta; (iii) within the genus Iberolacerta three groups were found: a Pyrenean group, an Iberian group and I. horvathi. The relationships among these groups remain unresolved; and (iv) the Peña de Francia lizards, described originally as a subspecies of I. cyreni, are in fact more closely related to I. monticola.

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Six nominal taxa (`Lacerta` monticola cantabrica, `L. ` monücola monticola, `L. ` aranica, `L. ` aurelioi, `L. ` bonnali, `L. ` cyreni) of the Iberian-Pyrenean Mountain Lizards were investigated by allozyme electropho- resis with regard to IS enzymatic loci. `L. ` horvathi was used as an outgroup for phylogenetic analysis. On the basts of our results - which are in perfect agreement with karyological data (ODIERNA & al. in press a, b) - five taxa can be distinguished (`£.`. monticola, `L`. aranica, `L`. aurelioi, `L`. bonnali, %`. cyreni) which we consider to be species.

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The albumins from 41 species of the family Lacertidae representing a substantial part of genera as well as infrageneric groups of the collective genus Lacerta s.l. were investigated by means of the MCF technique. The data allow a chronological estimation of the most important radiation events and their correlation with paleogeographical facts. Our results are in remarkable contradiction to the phylogeny presented by ARNOLD (1989a) based mainly on morphological characters. The most important phylogenetic relationships are: 1. The genera Gallona and Psammodromus are already isolated from the other taxa since the Oligocene. We assess these main branches as subfamilies Gallotiinae and Lacertinae. 2. Lacerta s.str. and Lacerta subgenus Zootoca form the sister group of the remaining Lacertinae. 3. A colonization of Africa in the Lower Miocene has led to a separation in two lines, with mainly Eurasian and African members respectively. 4. ARNOLD`S (I.e.) `Ethiopian and advanced Saharo-Eurasian clade` proved to be a paraphyletic unit, one group of genera being more closely related to European taxa. All techniques applied to the systematics of Lacertidae hitherto are critically discussed. An area-time- hypothesis concerning the phylogeny of Lacertidae is presented. Key words: Lacertidae, phylogeny of; Gallotiinae subfam. nov.; Lacertinae subfam. nov.; micro complement fixation; albumin evolution.

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The family Lacertidae encompasses more than 250 species distributed in the Palearctis, Ethiopis and Orientalis. Lacertids have been subjected in the past to several morphological and molecular studies to establish their phylogeny. However, the problems of convergent adaptation in morphology and of excessively variable molecular markers have hampered the establishment of well supported deeper phylogenetic relationships. Particularly the adaptations to xeric environments have often been used to establish a scenario for the origin and radiation of major lineages within lacertids. Here we present a molecular phylogenetic study based on two nuclear marker genes and representatives of 37 lacertid genera and distinct species groups (as in the case of the collective genus Lacerta). Roughly 1600 bp of the nuclear rag1 and c-mos genes were sequenced and analyzed. While the results provide good support to the hitherto suggested main subfamilies of Gallotiinae (Gallotia and Psammodromus), Eremiainae and Lacertinae [Harris, D.J., Arnold, E.N., Thomas, R.H., 1998. Relationships of lacertid lizards (Reptilia: Lacertidae) estimated from mitochondrial DNA sequences and morphology. Proc. R. Soc. Lond. B 265, 1939–1948], they also suggest unexpected relationships. In particular, the oriental genus Takydromus, previously considered the sister-group to the three subfamilies, is nested within Lacertinae. Moreover, the genera within the Eremiainae are further divided into two groups, roughly corresponding to their respective geographical distributions in the Ethiopian and the Saharo-Eurasian ranges. The results support an independent origin of adaptations to xeric conditions in different subfamilies. The relationships within the subfamily Lacertinae could not be resolved with the markers used. The species groups of the collective genus Lacerta show a bush-like topology in the inferred Bayesian tree, suggesting rapid radiation. The composition of the subfamilies Eremiainae and Lacertinae as well as their phylogeography are discussed.

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

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

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

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Infertile and non-viable fertile eggs within a reptile clutch may decrease the incubation success of the remaining eggs, as (1) opportunistic pathogens may use the nutrient resources provided by dead eggs to colonize the clutch and spread to and kill viable eggs; and (2) odours released by spoilt eggs may attract predators to the clutch. These hypotheses were tested on the Iberian rock lizard (Lacerta monticola) by comparing the incubation success of fertile eggs between clutches composed solely of fertile eggs and clutches containing a small number of dead eggs. In a laboratory experiment, fungi (Fusarium sp. and Gliocladium sp.) colonized both infertile eggs and fertile eggs that died during incubation and thereafter spread to and killed adjacent eggs. In addition, offspring hatched earlier from fungal infected eggs than from non-infected eggs. The former were smaller and lighter than the latter, as they hatched before using the full egg yolk content. Results from a field experiment did not corroborate the fungal pathogenic effects observed in the laboratory nor did they confirm that clutches containing dead eggs suffer higher predation. Despite the inconclusive results regarding the role of fungal pathogens in nature, the present study indicates that this subject deserves further investigation in reptiles.

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1. Iberian Rock Lizard ( Lacerta monticola , Boulenger 1905) males produce copulatory plugs that harden and adhere firmly inside the female cloaca immediately after copu- lation and occlude both oviductal openings. 2. To determine whether plugs reduce the chance of females being inseminated by rival males, two hypotheses were tested: (i) that plugs reduce female attractiveness and/or receptivity to rival males; and (ii) that plugs function as ‘chastity belts’. 3. There was no evidence to support either hypothesis. Neither male initiation of copulations and female termination of male copulation attempts, nor male insemina- tion success differed significantly between trials in which males were presented with females without a plug and with females with a plug deposited by a different male 1, 3 and 6 or more hours earlier, nor between females without and with a plug. 4. Several hypotheses for the function of Iberian Rock Lizard plugs are discussed. Since rival males were frequently able to displace a plug in the female cloaca, our results are consistent with a ‘sperm protection’ function of male postcopulatory mate guarding in squamate reptiles, but this remains to be tested.

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Male Iberian rock lizards (Lacerta monticola) produce copulatory plugs that adhere firmly inside the female cloaca and occlude both oviducts. These plugs do not prevent rival male insemination, as they neither reduce female attractiveness or receptivity to rival males nor do they function as “chastity belts”. Prior to copulation, males bite various regions of the female body, including the cloacal region. A previous plug in the female cloaca is expelled only after rival male intromission. We hypothesized that: (1) such male pre-copulatory behaviours function to loosen plug adherence to the female cloaca, thereby facilitating intromission, and that (2) the hemipenis plays a role in displacing a previous plug prior to the delivery of a new plug. Neither of these hypotheses was supported. Instead, our results indicate that rival males can intromit the hemipenis past a previous plug in the female cloaca and deliver their own plug underneath it. Consequently, previous plugs are pushed away from the oviductal openings and even dislodged from the female cloaca. Copulation duration was determined both by the time used by males to deliver a plug and by the fact that males prolonged copulation beyond plug delivery. There seemed to be sexual conflict over prolonged copulation, which was resolved by the male/female headlength ratio. The adaptive value of Iberian rock lizard copulatory plugs and prolonged copulation in Iberian rock lizards is discussed in the context of sperm competition.

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Socially dominant males often signal their status to rival males and/or females. We tested the hypotheses that Lacerta monticola femoral gland secretions and copulatory plugs convey chemical information about male identity and dominance status. We estimated male dominance status by staging male-male agonistic encounters in a neutral arena. We then conducted two experiments to compare male tongue-flick behavior toward chemical stimuli consisting of cotton swabs bearing (1) deionized water (control), the lizard`s own femoral secretions, and the femoral secretions of another male and (2) phosphate-buffered saline solution (control), the lizard`s own plug products, and the plug products of another male. Results indicate that males discriminated their own femoral secretions and plugs from those of other males. They also discriminated morphological attributes of other males that were associated with dominance status based on chemical cues arising from femoral secretions and discriminated the dominance status of other males based on chemical cues arising from the plugs. Femoral secretions that convey information about male identity and dominance status may be hypothesized to function in the establishment of L. monticola dominance hierarchies through scent-marking of territories. We suggest that copulatory plugs and femoral secretions may allow males to scent-mark the female body and postulate that this behavior may influence male and female reproductive decisions under selective pressures of sperm competition.

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Faeces play a role in intraspecific chemical communication in many vertebrates, including lizards. Here, we hypothesised that juvenile Iberian rock lizards (Lacerta (=Iberolacerta) monticola) use substrate-borne conspecific faecal chemicals to assess the social environment and base their spatial decisions. To test this hypothesis, we prepared chemical stimuli by dissolving faeces in an organic solvent (dichloromethane, DCM) and conducted chemosensory trials where we quantified latency to the first tongue flick (TF) and TF rates when young captive-borne juveniles were placed in an arena compartment whose substrate was labelled by (1) a control (DCM), (2) their own faecal chemicals, (3) faecal chemicals of another juvenile, (4) faecal chemicals of an adult female or (5) faecal chemicals of an adult male. Following TF observations, we removed a partition that separated the labelled compartment from a chemically unlabelled one, and recorded for 5 min when juveniles first crossed to the unlabelled compartment and total time spent in the labelled compartment. Each juvenile was tested with all stimuli in a randomised order. In addition, juveniles belonged to different families and were unrelated to and unfamiliar with faeces donors. Taken together, TF and spatial responses toward stimuli indicate that juveniles discriminated between faecal chemicals of conspecific juveniles, adult females and males, and that they avoided remaining in substrates labelled by adult male faecal chemicals. We suggest that juveniles assess the social environment based on conspecific faecal chemicals thus avoiding aggression and cannibalistic risks undertaken from encounters with adult males.

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Sperm competition selects for opposing male defensive and offensive reproductive traits, and its outcome may be determined by the effectiveness to which one trait has evolved to out-compete the other. We tested the effectiveness of a first male plug physical interference with a second male insemination (defence) vs the effectiveness of plug and associated sperm displacement by a second male (offence) on the outcome of sperm competition in Iberian rock lizards. We conducted a double mating experiment where we compared the proportion of eggs per clutch fertilised by the same second males (against the same first males) when they copulated with females 30 min (first male plug adhered firmly inside the female cloaca) and 4 h (first male plug loosely adhered or shed from the female cloaca) after first males. We found that second males fertilised the majority of the eggs per clutch in the 30-min treatment, whereas fertilisations were equally shared between the two males in the 4-h treatment. These results show that plugs have little defensive effectiveness, and thus, do not assure high first male fertilisation success. Instead, sperm displacement appears to be associated with plug displacement. That is, because sperm embedded in first male plugs, and displaced from competition for fertilisations by second males, is expected to increase in number with decreasing time allowed for female sperm transport, second males thus enjoy higher fertilisation success. This study shows that offensive plug displacement out-competes plug defensive role in Iberian rock lizards. Moreover, it reveals sperm displacement as a novel sperm competition mechanism in reptiles.

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A BLACK SPECIMEN OF LACERTA MONT/COLA Isolation and environmental factors can result in special colored populations of lizards on moun- taintops and islands. Near Lago Ercina in the Pi- cos d`Europa a single all black specimen ofLacer- ta monticola was found. Although this specimen could be infertile, ill or not capable of getting a suitable territorium, the author nevertheless sug- gests the possibility of the development of a black population in about ten years time.

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We conducted a karyological study of Iberian rock lizards by standard and C-, AgNOR-, DAPI-, and Alu I-banding methods. The results showed that Lacerta monticola monticola and L. m. cantabrica possess 2N = 36 uniarmed chromosomes. Lacerta m. cyreni differs from the other two subspecies of L. monticola in the NOR-bearing chromosome pairs and in the presence of the heteromorphic and heterochromatic W-chromosome. The Pyrenean species L. aurelioi and L. bonnali differ from L. monticola in having a karyotype rich in biarmed elements. In fact, L. aurelioi possesses a 2N = 26 karyotype with 10 biarmed + 16 uniarmed macrochromosomes. In L. bonnali, an intraspecific variability in diploid number due to Robertsonian translocations was found: the specimens of L. b. aranica have a karyotype similar to that of L. aurelioi while the specimens from nominal subspecies (population of Monte Perdido) possess a 2N = 24 karyotype with 12 biarmed + 12 uniarmed macrochromosomes. Our results confirm that Iberian rock lizards form a heterogeneous group in which at least five taxa can be distinguished karyologically.

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The occurrence and form of sex chromosomes were investigated with the aid of C-banding and 4′-6-diamidino-2-phenylindole (DAPI) staining in 13 species of lacertid lizards. The results obtained show the presence in five species of a female heterogamety in which the two sex chromosomes have the same shape and size, but the W differs from the Z in being almost entirely heterochromatic. This condition is clearly similar to that found in some snakes and considered to be an early stage of differentiation of sex chromosomes by Singh et al. (1976, 1980). A more evolved condition may be that found in three other species in which the W is distinctly smaller than the Z. A third situation is that found in all Podarcis species which, even though they are considered to be among the more evolved species in the family, possess two sex chromosomes that are indistinguishable. In general, the situation in lacertids may be compatible with the hypothesis of sex chromosome evolution put forward by Singh et al. (1976, 1980). However a differentiation mechanism of this kind does not seem to be well established in lacertids, and is probably not the only mechanism that is in operation in this family.

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We studied the thermal ecology of the montane Iberian rock lizard, Iberolacerta monticola, in the western area of its distribution at the Serra da Estrela (Portugal). We calculated the precision of thermoregulation and the indices of thermal quality of the habitat, and accuracy and effectiveness of thermoregulation. To complete the study of the thermal ecology, we assessed the relationships between body and environmental temperatures, and we described the thermal and spatial heterogeneity of the habitat. Our results indicate that the Iberian rock lizard is a cold-specialist, with a preferred temperature range between 29.80 and 31.60 °C. Thus, precision of thermoregulation is 1.8 °C, which is a normal range in thermal specialists, like other species of the genus Iberolacerta. This result is important because being thermal specialists and living in mountaintops make Iberian mountain lizards particularly vulnerable to global warming. The habitat of I. monticola at the Serra da Estrela is formed of microhabitats offering different operative temperatures, which allows lizards to select the most suitable for thermoregulation at any time of the day. Iberian rock lizards achieve an effectiveness of thermoregulation of 0.86, thanks to careful thermoregulatory behaviour. Rocky microhabitats occupy more than 50% of its habitat, so is probable that lizards are selecting rocks to warm themselves faster, minimising the costs of thermoregulation. A possible thigmothermic component of this kind would be unique among the species of Iberolacerta.

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Lacertinae is one of the three lacertid lizard subfamilies with a geographical distribution confined to the Palaearctic. Several past attempts to reconstruct its phylogeny resulted in unresolved bush-like topologies. We address the question of whether the lack of resolution is due to insufficient data or whether this lack reflects a rapid succession of speciation events. We analyzed four partial and one complete gene sequences from mitochondrial and nuclear genomes, totalling roughly 3600 bp. We included 29 species representing all 19 genera suggested in recent revision of Lacertinae [Arnold, E.N., Arribas, O., Carranza, S., 2007. Systematics of the palaearctic and oriental lizard tribe Lacertini (Squamata: Lacertidae: Lacertinae), with descriptions of eight new genera. Zootaxa 1430, 1–86]. The resulting phylogeny, first, corroborates monophyly at the genus level for the suggested genera, as well as the finding that Atlantolacerta andreanskyi, until recently part of Lacertinae, belongs to the subfamily Eremiadinae. Second, we find that increasing the sequence length and combining multiple nuclear and mitochondrial sequences did not resolve the polytomy, suggesting that the inferred topology indicates a multiple cladogenesis within a short geological period, rather than a methodical artefact. Divergence time estimates, based on previous estimates of several node ages, range from 13.9 to 14.9 million years for the radiation event, however with very broad confidence interval. To associate the radiation with a narrower geological time we consider palaeogeographic and palaeoclimatic data, assuming that the Lacertinae probably evolved in Central Europe andWAsia after the collision of Africa and Eurasia. We suggest that this radiation may date to the late Langhian (ca. 14–13.5 million years) when geological events caused abrupt changes in regional water–land distribution and climate, offering a window of distinct conditions.

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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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We studied diet composition and prey selection in the lizard Lacerta monticola throughout its activity season. The most important prey groups in terms of numbers were Coleoptera, Diptera, Formicidae, and Araneae. Comparison of diet composition in the adult lizards with estimates of prey availability indicated that lizards exhibited precisely defined patterns of prey selection. Electivities were negative for prey < 3 mm in length and generally positive for larger prey, despite the higher relative abundance of the former group in the environment. Electivity scores for individual prey taxa were positively correlated with their length, and lizards preferentially consumed larger individuals belonging to the taxa containing smaller animals. Relative availability of the four major prey taxa fluctuated considerably throughout the year. In spite of these changes, monthly rates of consumption of Araneae and Diptera remained almost constant. Variations in monthly electivity scores were negatively correlated with relative availability of three (Coleoptera, Diptera, and Araneae) of the four major prey taxa. These results fit the predictions of prey-selection models based on nutrient constraints.

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

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When prey take refuge to reduce predation risk, they forfeit time for other activities. They may also pay a physiological cost. In particular, optimal regulation of body temperature is essential for ectotherms. Qualitative models predict that lizards have to balance anti-predatory decisions in relation to thermal conditions of the refuge and predation risk when deciding when to resume activity. However, these models are not general and empirical tests of their assumptions are scarce. We modified previous models to include the case of a high and maintained level of predation risk. The predictions of the model were further investigated in a laboratory experiment using male Iberian rock lizards. The same level of predation risk was displayed in two treatments in which temperature inside the refuge was high or low, and in the mating or the post-reproductive season. As predicted, lizards increased successive emergence times - an increase that was not linear but accelerating - and they had shorter emergence times when thermal costs of refuge use were higher. Nevertheless, body size of lizards and the season of the experiment had no effect on these decision rules. Our results are in line with an economical balance between costs and benefits in the decision rules controlling active versus inactive periods

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

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The fragmentation and destruction of natural habitats by human intervention is producing a continuous and inexorable reduction of the size of populations in multitude of species all over the world. Small and isolated populations face higher extinction risks, due to demographic and environmental stochasticity, and also because of several genetic threats, among which inbreeding is considered the most important one. For many of these species, the extinction of a population is an irreversible event, so that determining the immediate importance of these risk factors and understanding their interactions is crucial for conservation plans. Iberolacerta monticola is a small lacertid endemic to the northwestern Iberian Peninsula, distributed mainly across moderate/high altitude mountainous regions. Some populations are found nearly at sea-level, though, in fluvial valleys with relict Atlantic forests, in the severely fragmented western part of its range. One of them has been dramatically reduced over the last 30 years, and presently is on the brink of extinction. Using microsatellite nuclear markers, we obtained different measures of genetic variation at this site, together with demographic and breeding data. Both the level of heterozygosity and the number of alleles per locus indicate that the level of variation in this population is comparatively high, and the average inbreeding coefficient is very low. Individuals appear healthy and long-lived, and are related by a few different lines of descent. These findings are discussed in the context of current theories and experimental evidence of associative overdominance and purging of the genetic load of populations, with special emphasis on the evolutionary potential of recovery of small evolutionary units.

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Aim The study of the factors that influence population connectivity and spatial distribution of genetic variation is crucial for understanding speciation and for predicting the effects of landscape modification and habitat fragmentation, which are considered severe threats to global biodiversity. This dual perspective is obtained from analyses of subalpine mountain species, whose present distribution may have been shaped both by cyclical climate changes over ice ages and anthropogenic perturbations of their habitats. Here, we examine the phylogeography, population structure and genetic diversity of the lacertid lizard Iberolacerta monticola, an endemism considered to be facing a high risk of extinction in several populations. Location Northwestern quadrant of the Iberian Peninsula. Methods We analyzed the mtDNA variation at the control region (454 bp) and the cytochrome b (598 bp) loci, as well as at 10 nuclear microsatellite loci from 17 populations throughout the distribution range of the species. Results According to nuclear markers, most sampling sites are defined as distinct, genetically differentiated populations, and many of them show traces of recent bottlenecks. Mitochondrial data identify a relatively old, geographically restricted lineage, and four to six younger geographically vicariant sister clades, whose origin may be traced back to the mid-Pleistocene revolution, with several subclades possibly associated to the mid-Bruhnes transition. Geographic range fragmentation of one of these clades, which includes lowland sites, is very recent, and most likely due to the accelerated loss of Atlantic forests by human intervention. Main Conclusion Altogether, the data fit a “refugia within refugia” model, some lack of pattern uniformity notwithstanding, and suggest that these mountains might be the cradles of new species of Iberolacerta. However, the changes operated during the Holocene severely compromise the long-term survival of those genetic lineages more exposed to the anthropogenic perturbations of their habitats.

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Fourteen polymorphic microsatellite loci are described for the Iberian rock lizard, Iberol- acerta monticola. Genetic variation in a sample of 20 individuals from Piornedo (north- western Spain) was quantified both by the number of alleles per locus, which ranged from six to 13, and by the expected frequency of heterozygotes under random mating (heterozy- gosity), which ranged from 0.761 to 0.902. Single locus and global exclusion probabilities were also computed, and indicate a high power of these markers for paternity assignments and mating system studies of I. monticola. All the analysed loci were also polymorphic in Iberolacerta galani, but only seven in Zootoca vivipara.

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Rock lizards of the genus Iberolacerta constitute a promising model to examine the process of sex chromosome evolution, as these closely related taxa exhibit remarkable diversity in the degree of sex chromosome differentiation with no clear phylogenetic segregation, ranging from cryptic to highly heteromorphic ZW chromosomes and even multiple chromosome systems (Z1Z1Z2Z2/Z1Z2W). To gain a deeper insight into the patterns of karyotype and sex chromosome evolution, we performed a cytogenetic analysis based on conventional staining, banding techniques and fluorescence in situ hybridization in the species I. monticola, for which previous cytogenetic investigations did not detect differentiated sex chromosomes. The karyotype is composed of 2n = 36 acrocentric chromosomes. NORs and the major ribosomal genes were located in the subtelomeric region of chromosome pair 6. Hybridization signals of the telomeric sequences (TTAGGG)n were visualized at the telomeres of all chromosomes and interstitially in 5 chromosome pairs. C-banding showed constitutive heterochromatin at the centromeres of all chromosomes, as well as clear pericentromeric and light telomeric C-bands in several chromosome pairs. These results highlight some chromosomal markers which can be useful to identify species-specific diagnostic characters, although they may not accurately reflect the phylogenetic relationships among the taxa. In addition, C-banding revealed the presence of a heteromorphic ZW sex chromosome pair, where W is smaller than Z and almost completely heterochromatic. This finding sheds light on sex chromosome evolution in the genus Iberolacerta and suggests that further comparative cytogenetic analyses are needed to understand the processes underlying the origin, differentiation and plasticity of sex chromosome systems in lacertid lizards.

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Satellite DNAs compose a large portion of all higher eukaryotic genomes. The turnover of these highly repetitive sequences is an important element in genome organization and evolution. However, information about the structure and dynamics of reptilian satellite DNA is still scarce. Two satellite DNA families, HindIII and TaqI, have been previously characterized in four species of the genus Iberolacerta. These families showed different chromosomal locations, abundances, and evolutionary rates. Here, we extend the study of both satellite DNAs (satDNAs) to the remaining Iberolacerta species, with the aim to investigate the patterns of variability and factors influencing the evolution of these repetitive sequences. Our results revealed disparate patterns but also common traits in the evolutionary histories of these satellite families: (i) each satellite DNA is made up of a library of monomer variants or subfamilies shared by related species; (ii) species-specific profiles of satellite repeats are shaped by expansions and/or contractions of different variants from the library; (iii) different turnover rates, even among closely related species, result in great differences in overall sequence homogeneity and in concerted or non-concerted evolution patterns, which may not reflect the phylogenetic relationships among taxa. Contrasting turnover rates are possibly related to genomic constraints such as karyotype architecture and the interspersed organization of diverging repeat variants in satellite arrays. Moreover, rapid changes in copy number, especially in the centromeric HindIII satDNA, may have been associated with chromosomal rearrangements and even contributed to speciation within Iberolacerta.

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Telomeres are nucleoprotein complexes protecting the physical ends of linear eukaryotic chromosomes and therefore helping to ensure their stability and integrity. Additionally, telomeric sequences can be localized in non-terminal regions of chromosomes, forming so-called interstitial telomeric sequences (ITSs). ITSs are traditionally considered to be relics of chromosomal rearrangements and thus very informative in the reconstruction of the evolutionary history of karyotype formation. We examined the distribution of the telomeric motifs (TTAGGG)n using fluorescence in situ hybridization (FISH) in 30 species, representing 17 families of squamate reptiles, and compared them with the collected data from another 38 species from literature. Out of the 68 squamate species analyzed, 35 possess ITSs in pericentromeric regions, centromeric regions and/or within chromosome arms. We conclude that the occurrence of ITSs is rather common in squamates, despite their generally conserved karyotypes, suggesting frequent and independent cryptic chromosomal rearrangements in this vertebrate group.

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We studied the reproductive ecology of a population of the lacertid lizard Lacerta monticola at a lowland location in A Coruña (NW Spain) from 1997 to 2002. The timing of the reproductive cycle was examined based on mark-recapture records of individual lizards in the Ž eld. The characteristics of the eggs and hatchlings were obtained from clutches laid in the laboratory by pregnant females that were temporarily removed from the study area. Our results indicate that mating took place between late March and July. The smallest female with signs of sexualmaturity had a snoutvent length (SVL) of 52.4 mm and the smallest male was 50.7 mm. Sexual maturity was attained at the age of 2 years by 41% of the individuals, while the remaining 59% of the lizards matured at the age of 3 years. The laying period occurred between June and the beginning of August considering all years. About 61% of the reproductive females produced a single clutch annually, while 39% of the females produced two clutches per year. Females that produced two clutches were generally larger (average SVL D 70.9 mm) than those that laid a single annual clutch (average SVL D 62.6 mm). The mean clutch size was 6.4 eggs (range 4-9). Both clutch size and clutch mass increased signiŽ cantly with female SVL. The mean egg mass in a clutch decreased signiŽ cantly with clutch size. Hatching occurred between August and September. Hatching success in the laboratory was 71.7%. The mean SVL was higher in female hatchlings than in males.

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

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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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Elements of the herpetofauna are in the author`s opinion of great importance for the zoogeographical division of the Palaearctic region. It is suggested that this region be divided into the following four subregions: 1/ arctic, 2/ boreal Euro-Siberian, 3/ bo- real Himalayan-Manchurian and 4/ Mediterranean-Central-Asiatic.

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Evolutionary changes in foraging style are often believed to require concurrent changes in a complex suite of morphological, physiological, behavioural and life-history traits. In lizards, species from families with a predominantly sit-and-wait foraging style tend to be more stocky and robust, with larger heads and mouths than species belonging to actively foraging families. Here, we test whether morphology and foraging behaviour show similar patterns of association within the family Lacertidae. We also examine the association of bite force abilities with morphology and foraging behaviour. Lacertid lizards exhibit considerable interspecific variation in foraging indices, and we found some evidence for a covariation between foraging style and body shape. However, the observed relationships are not always in line with the predictions. Also, the significance of the relationships varies with the evolutionary model used. Our results challenge the idea that foraging style is evolutionarily conservative and invariably associated with particular morphologies. It appears that the flexibility of foraging mode and its morphological correlates varies among lizard taxa.

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Apart from certain species of the African clade, the lizard family Lacertidae has generally been described as consisting of active foragers. We made quantitative field observations of 14 species of lacertid lizards, mainly belonging to the more basal Eurasian clade. Our data show that sit-and-wait foraging is much more widespread in Lacertidae than previously alleged. We also investigated the influence of weather and times of day on foraging activity levels and conclude that for comparative purposes observations should be restricted to circumstances that are optimal for activity. We did not find sex differences in foraging behavior.

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Foraging strategy is often considered to play a central role in moulding diverse aspects of an animal`s general biology. Active foragers should have greater locomotor endurance, allowing high movement activity rates, while sit-and-waiting foragers may be better adapted to sprinting, allowing catching prey by a quick attack from an ambush site, and going with specific predator escape tactics. In this study we investigate these predicted patterns in a set of lacertid lizard species. There is considerable variation in foraging activity within Lacertidae, which allows the close investigation of the co-evolution of the traits considered. We found a tendency of positive correlation of foraging measures (PTM, percentage of time moving; MPM, number of movements per minute) with laboratory measured endurance capacity. However, the relationship of foraging measures with maximal sprint speed remains less clear. MPM correlates negatively with maximal sprint speed, but PTM does not. When sprint speed was corrected for body size, no correlations were found at all.

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Space usage by animals may be influenced by a range of factors. In this study we investigate whether foraging behaviour affects the home range size of lizards. Two distinct tactics of foraging have been recognized in predators: sit-and-wait foraging (SW) and active foraging (AF). Foraging activity level of a data set of lizard species, mainly compiled from literature, is compared with their home range sizes. Two opposite predictions can be made about foraging in connection with home range area: on the one hand, SW species may exhibit larger home ranges due to their mating system; on the other hand, AF species have higher metabolic energy and thus food requirements and can be expected to have larger home ranges that have to yield this food. This study shows that percentage of the time moving (as an index of foraging mode) correlates positively with home range, even after correcting for body mass, and these patterns remain when phylogenetic relationships are taken into account. We thus conclude that home range areas parallel activity levels in lizards.

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Environmental contamination is supposed to be a reason for population declines in reptiles. Especially intensification and expansion of agriculture are leading to increased pesticide exposure risks for wildlife. In the European Union, Special Areas of Conservation (SACs) have been established for the conservation of taxa listed in Annex II of the Habitats Directive. In the SACs, agricultural land use is legal. Therefore, we conducted a risk evaluation of pesticide exposure for Annex II reptiles by calculating proportions of land use with regular pesticide applications within SACs. Using three evaluation factors (occurrence probability, physiology, life-history aspects), a species-specific risk index was created. Nearly half of the species at above-average risk by pesticide use are globally threatened with extinction (IUCN Red List of Threatened Species). About 30% of their SACs are agriculturally used and one priority subspecies of the Habitats Directive is at highest risk (Vipera ursinii rakosiensis). Also, all evaluated fresh-water and land-dwelling turtle species are at high risk. National variation in agricultural land use in the SACs was observed. Species at above-average risk are mainly distributed in the Mediterranean and Pannonian/Continental biogeographical regions of Europe. Conservation status according to the IUCN Red List of Threatened Species as well as national differences among the member states argue for the inclusion of pesticide risk assessments in site-specific management plans for SACs to avoid regional loss of reptilian biodiversity.