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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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Thermal constraints may limit the physiology and behaviour of ectotherms because of the high thermal dependence of metabolic functions. The adaptive mechanisms of thermoregulation and the cost of confronting thermal constraints were studied in the Cyren’s Rock Lizard (Iberolacerta cyreni (Muller and Hellmich, 1937)), a lacertid lizard endemic to mountain areas of central Spain. Semicontinuous monitoring of body temperature (Tb) in the laboratory indicated that the preferred temperature range for this population (Tpref) was lower than those found for most lacertid lizards, and field body temperatures of active animals in summer were even lower than Tpref. Overall these results, together with distribution of field operative temperatures (Te), indicate that I. cyreni is an active and relatively accurate thermoregulator, although limited by thermal constraints in their habitat. Laboratory experiments in contrasting thermal environments showed that even under thermally restricted conditions, lizards achieved their Tpref by modifying their thermoregulatory behaviour, principally through changes in space use, basking time, and body posture. However, these behavioural adjustments to reach the Tpref have associated costs, and lizards spent 80% of their time in thermoregulation when tested under low radiation conditions, which in the wild would limit the scope for other activities and eventually increase predation risk. Our results suggest that thermoregulatory behaviour may play an important role in coping with global climate change, hence predictions of the effects of climate warming on lizards inhabiting cold habitats should take into account the buffering role of behavioural thermoregulation.

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We analysed asymmetry in the common wall lizard (Podarcis muralis) and the Horvath`s rock lizard (Iberolacerta horvathi) from 16 populations in Slovenia and Croatia to understand their connection with potential stress factors: altitude, urbanization and the presence of another species (interspecific competition). We also compared morphological body characteristics between the two species. We used geometric morphometric methods to create landmark coordinates in the photographs of lizards` heads. Modified ANOVAs and an asymmetry index were used to find the differences in four physical characteristics: shape and size of the head, body size and the number of supraciliar scales. We found fluctuating asymmetry in the number of supraciliar scales and the shape of the head in all of our samples. Both species had more asymmetric heads in the intermediate altitudes, which might be connected with lower availability of suitable habitats. Common wall lizards from urban environment did not have more asymmetries compared to those living in natural environments. Contrary to our expectations, we found more asymmetries in allotopic than syntopic populations, indicating limited effect of interspecific interactions. Horvath`s rock lizards had more asymmetric heads, which suggests they might be exposed to greater environmental stress, although genetic effects are also possible, since directional asymmetry was also detected.

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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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We report the first study of diet composition of a Pyrenean lizard of the genus Iberolacerta, the Aurelio`s lizard, I. aurelioi, living in an alpine rocky slope at 2300 m of altitude in Andorra. Diet composition was studied during two years from 289 faecal pellets containing 966 prey items. Number of prey per pellet shows annual, seasonal and sexual differences. Diet of the species is poorly diverse, mainly dominated by insects, but we find clear evidences of opportunistic cannibalism based on the presence of toes and scales in feces. In addition, we detected minor sexual, annual and month differences in prey presence and abundance. Low amplitude of trophic niche width and prey diversity, as well as predation on clumped prey and cannibalistic behaviour suggest that Iberolacerta aurelioi has a remarkable dietary strategy in response to strong thermal and food availability constraints at high mountain habitats.

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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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We analyzed whether habitat deterioration due to ski slopes affected lizards’ behavior, and whether these changes in behavior had consequences for the body condition and health state of lizards. Results suggested that habitat deterioration in ski slopes not only implied a loss of optimal habitat for lizards, but also led to an increase in perceived risk of predation. Males seemed to adjust their movement patterns to differences in risk, increasing their movement speed during their displacements across risky areas within ski slopes, but as a consequence, they incurred loss of body condition. A laboratory experiment supported that fleeing at high speeds in areas without refuges can be a factor responsible for reduced body condition. However, changes in body condition did not affect sprint speed of lizards. Our study provides new evidence that behavioral strategies to cope with increased predation risk, due to human-induced habitat deterioration, may affect body condition of lizards. Our results have applications for the design of conservation plans for this endangered lizard species. Environmental impact studies should be performed before opening new ski slopes, but taking into account not only the presence of lizards but also their body condition. An effective way to decrease the effects of habitat loss may be the artificial restoration of some refuges to create safe corridors for movement.

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Lizards often respond to increased predation risk by increasing refuge use, but this strategy may entail a loss of thermoregulatory opportunities, which may lead to a loss of body condition. This may be especially important for pregnant oviparous female lizards, because they need to maintain optimal body temperatures as long as possible to maximize developmental embryos rate until laying. However, little is known about how increased time spent at low temperatures in refuges affects body condition and health state of pregnant female lizards. Furthermore, it is not clear how initial body condition affects refuge use. Female Iberian rock lizards forced to increase time spent at low temperatures showed lower body condition and tended to show lower cell-mediated immune responses than control females. Therefore, the loss of thermoregulatory opportunities seems to be an important cost for pregnant females. Nevertheless, thereafter, when we simulated two repeated predatory attacks, females modified refuge use in relation to their body condition, with females with worse condition decreasing time hidden after attacks. In conclusion, female lizards seemed able to compensate increased predation risk with flexible antipredatory strategies, thus minimizing costs for body condition and health state.

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Prey often respond to predation risk by increasing refuge use, but this strategy may entail a loss of body condition. Factors responsible for this loss of body condition remain unclear. Also how prey deal with refuge use to cope with predation risk without incurring costs of body condition, and how initial body condition affects refuge use remain barely known. We analyzed in the field whether adult Iberolacerta cyreni lizards modify their escape strategies and refuge use in areas with different levels of habitat deterioration and ecotourism pressure, which represent different levels of predation risk, and the consequences of changes in antipredator behavior to body condition. Lizards inhabiting deteriorated areas, where risk is higher, remained closer to refuges, but decreased time spent hidden in refuges after attacks, probably to maintain similar body condition than lizards inhabiting natural areas. We performed two laboratory experiments to isolate potential costs of refuge use that might affect the body condition of male lizards: a) a decrease of the efficiency of digestion due to low temperatures inside refuges and/or b) a reduction in food intake. Results suggest that refuge use is costly in terms of body condition due to reduced food intake, but the loss of efficiency of digestion seems unimportant. Lizards modified refuge use in relation to their body condition, with lizards with worse condition decreasing time hidden after predatory attacks. We concluded that lizards compensated for increased predation risk with flexible antipredatory strategies, coping with risk without incurring costs for body condition.

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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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Variation in environmental factors plays a central role on organisms` physiological changes. However, the physiological response to predation risk has rarely been investigated in reptiles. Chemical senses are important for intraspecific communication in squamate reptiles. In male lizards Iberolacerta cyreni the maintenance of relative proportions of lipids in femoral gland secretions is costly, which may ensure honest signalling of quality. We hypothesized that increased predation risk should compromise the maintenance of such lipid proportions, as both a fear response and escaping behavior can have physiological consequences. We simulated predator attacks and found that relative proportions of lipids in femoral gland secretions changed in disturbed lizards but not in control ones. Thus, predator–prey interactions may modulate relative concentrations of chemicals in scents of lizards. Potential consequences of this effect on intraspecific chemical communication are suggested.

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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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Differences in surface structure (ober- hautchen) of body scales of lacertid lizards involve cell size, shape and surface profile, presence or absence of fine pitting, form of cell margins, and the occurrence of longitudinal ridges and pustular projections. Phylogenetic information indicates that the primitive pattern involved narrow strap-shaped cells, with low posteriorly overlapping edges and relatively smooth surfaces. Deviations from this condition produce a more sculptured surface and have developed many times, although subsequent overt reversals are uncommon. Like variations in scale shape, different patterns of dorsal body microornamentation appear to confer different and conflicting performance advantages. The primitive pattern may reduce friction during locomotion and also enhances dirt shedding, especially in ground-dwelling forms from moist habitats. However, this smooth microornamentation generates shine that may compromise cryptic coloration, especially when scales are large. Many derived features show correlation with such large scales and appear to suppress shine. They occur most frequently in forms from dry habitats or forms that climb in vegetation away from the ground, situations where dirt adhesion is less of a problem. Microornamentation differences involving other parts of the body and other squamate groups tend to corroborate this functional interpretation. Microornamentation features can develop on lineages in different orders and appear to act additively in reducing shine. In some cases different combinations may be optimal solutions in particular environments, but lineage effects, such as limited reversibility and different developmental proclivities, may also be important in their genesis. The fine pits often found on cell surfaces are unconnected with shine reduction, as they are smaller than the wavelengths of most visible light.

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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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Lacerta monticola bonnali Lantz, 1927, that inhabits central Pyrenees from 1.800 m to ca. 3.000 m a.s.l. in alpine and subalpine environments, should be considered hereinafter as a full species: L. bonnali stat. nov. due to strong differences in pholidosis, design and coloration, as well as ecological (types of rocks among it lives) and biological features (egg incubation time) that distinguish it both from L. monticola Boulenger, 1905 and L. horvathi Mehely, 1904, their closest geographic relatives, as well as from other Archeolacerta from Europe and Near East.

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Based on the study of geographic variability (pholidosis and biometry) of the Pyrenean endemite Lac bonnali LANTZ, 1927, a new subspecies of this lizard is described from Vali d`Aràn and Pallars.

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We give an historical review of the discovery and knowledge of A. bonnali in the Pyrenees and Catalonia. The up till now known locatities, habitat and specimens morphology are included.

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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 geographic limits of the three high mountain pyrenean lizards (Archaeolacerta bonnali, A. aranica and A. aurelioi) are relatively congruent with the biogeographic sectorizations proposed up till now, although don’t fit well with the hierarchy of these subdivisions. The differentiation degree among conspecific populations is significatively correlated with their separation distances. Our sectorization from the Alpine belt developed in basis to the alpine carabofauna is the first model specifically proposed for these biota. It clusters the alpine massifs in four groups: a) Occidental Pyrenees; b) Central Pyrenees (with a central portion s. str., an aranese portion, and a preoriental portion); and c) Eastern Pyrenees.

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Die Pyrenäenarten von Archaeolacerta s.l. (A. aranica, A. aurelioi und A. bonnali) unterscheiden sich von den übrigen Arten der Eurasiatischen Radiationslinie (sensu MAYER & BENYR 1994) durch den Besitz knöcherner Rippen im Bereich des dritten Präsakralwirbels. Die drei Pyrenäenarten zeigen in der Morphologie von Postorbitale und Postfrontale jeweils osteologische Verschiedenheiten, die Artniveau erreichen und ihre zweifelsfreie Unterscheidung erlauben. Auch finden sich Unterscheidungsmerkmale im Schultergürtel (Klavikel und Sternalfontanella). Apomorphe Merkmale innerhalb der kaukasischen ´Lacerta` saxicola – Gruppe stellen deutliche Unterschiede sowohl gegenüber den Archaeolacerten Europas (fast immer einzusätzlicher Präsakralwirbel) als auch gegenüber Apathya (ein zusätzlicher Präsakralwirbel und eine verminderte Anzahl von Prämaxillarzähnen) dar. ´L.` chlorogaster unterscheidet sich von den übrigen kaukasischen Formen klar durch die Zahl der Prämaxillarzähne und Form der Interklavikel. Die parthenogenetischen Taxa zeigen deutliche Ähnlichkeiten (Modalwerte der Wirbelzahl, Morphologie des Postorbitale) zu ihrem Elternarten, besonders zur weiblichen Elternart. Aufgrund der osteologischen Befunde wird deutlich, daß die europäischen, vorläufig in der Sammelgattung Archaeolacerta s.l. vereinigten Taxa keine natürliche Einheit darstellen. Die Formen der Pyrenäen und des Kaukasus erscheinen osteologisch deutlich differenziert, während die iberischen, balkanischen und tyrrhenischen Taxa in den untersuchten Strukturen nur synplesiomorphe Merkmalsausprägungen gemeinsam haben, die keine Aussagen über ihre Verwandtschaft ermöglichen.

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Auf Grundlage allen verfügbaren Materials werden neben einer umfangreiche Diagnose (Biometrie, Pholidose, Osteology, Karyotyp, Allozymdifferenzen, hemipenismorphologie) Angaben zu Verbreitung, Sexualdimorphismus und geographischer Variabilität von ´Lacerta` aurelioi ARRIBAS, 1994 vorgelegt. Pholidotische und metrische Merkmale werden dabei mit uni- und multivariatenn statistischen Methoden untersucht. `Lacerta` aurelioi ist in einem kleinen Bereich der östlichen Zentralpyrenäen endemisch. Die Art bewohnt die spanischen Gebirgsmassive Montroig und Pica d´Estats i. w. S. (inklusive Serrat de Capifonts) sowie NW-Andorra (Gebirgsstöcke Coma Pedrosa und Tristaina), woe sie zwischen 2100 m bis fast 2940 m Höhe gefunden wird. Dort kommt die Eidechse hauptsächlich auf kambrio-ordovizischen quarzitischen Schiefern und Phylliten, aber auch auf schwach metamorphosierten Staurolitschichten und schiefrigem Cordierit und Andalusit vor. Morphologisch zeiegen die untersuchten Populationen nur schwache Unterschiede auf infrasubspezifischem Niveau. Auch lassen sich keine Korrelationen zwischen Differenzierungsgrad und geographischer Lage oder Entfernung der Populationen feststellen: Die Subpopulationen eines gebirgsstockes (verschiedener benachbarter Hänge) sind häufig untereinander stärker verschieden als gegenüber Subpopulationen anderer Gebirgsstöcke, sodaß geographisch weiter voneinander entfernte Populationen einander morphologisch stärker ähneln können als geographisch nahe Populationen. Ein denkbares Szenario der Artbildung, das pleistozene Überleben und die Ausbreitung sowie die gegenwärtige Verbreitung der Art werden diskutiert. Die heutige Verschiedenheit der Populationen wird auf ihre holozäne Isolation zurückgeführt. Die Südhänge des Montroig Massives werden als ein würmzeitliches Refugium angesehen, von dem aus die Art ihr gegenwärtiges Verbreitungsgebiet, in dem die verschiedenen Populationen isoliert wurden, besiedelte.

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Iberolacerta bonnali is endemic from the Central Pyrenees, where inhabits from the Arriel Massif in the west, to the Aigüestortes mountains in the east. His range is characterized by the greater size of the mountain massifs which inhabits, and by to present granitic (igneous) rock substrates. By other side, his localities are also characterized by lesser insolations and sun radiations, as corresponds to the pyrenaean parts with greater atlantic influence. I. aranica is endemic from the Mauberme Massif and its spurs. His area appears characterized by greater snow accumulations both in winter and spring. I. aurelioi is endemic from the Montroig, Pica d`Estats and Coma Pedrosa massifs. His area is characterized by the greater insolations and sun radiations, and by the presence of lesser alpine belt surface and absolute heights of the inhabited massifs. Also, he inhabits only paleozoic rocks (mainly cambroordovician quarzites). In respect to other species present in the alpine belt: Lacerta agilis appears in localities characterized by high insolation and sun radiations in respect to other pyrenaean lizards. Zootoca vivipara do not present specific traits as lives in all the pyrenaean range and frequently sympatrically with all the other species here studied. Podarcis muralis appears in the comparatively more outer parts, with greater insolations, from the Pyrenees.

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

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Die Peña de Francia Gebirgseidechse (´L.` cyreni martinezricai) unterscheidet sich von anderen Formen ihrer Art (Gredos: ´L.` cyreni castiliana, Guadarrama: ´L.` cyreni cyreni) durch die signifikant verminderte Häufigkeit des Auftretens eines Schuppenkontaktes zwischen Rostrale und Internasale. Weiters zeichnen sich Männchen und Weibchen des Peña de Francia Taxons gewöhnlich durch blaue Ozellen in der Schultergegend und braune Rückenfärbung aus, während ´L.` cyreni anderer Populationen keine Ozellen tragen und sehr häufig grünrückig sind. Neben diesen sehr auffälligen (und evolutiv bedeutsamen) Farbmerkmalen finden sich signifikante aber nicht sehr starke Pholidoseunterschiede (Anzahl Rückenschuppen-Längsreihen, Bauchschuppen-Querreihen, Circumanalschilder). Das Vorkommen von ´L.` cyreni martinezricai ist auf die Gipfelregion eines isolerten Berges begrenzt. Die Populationsgröße (~ 50 adulte Individuen) nimmt ständig ab. Schutzmaßnahmen sind dringend erforderlich.

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Iberolacerta bonnali (LANTZ, 1927), ein Endemit der axialen Zentralpyrenäen, bewohnt die Gebirgsstöcke Arriel, Balaitous, Vignemale (oder Comachibosa), Panticosa, Monte Perdido, Punta Suelza,Posets, Maladeta, Ballibierna und die Gebirge des Sant Maurici-Aigüestortes Nationalparks (Besiberris, Muntanyó de Llacs, Peguera und Encantats) samt einer kleinen nördlichen Exklave in den Massiven von Neouvielle und Bigorre. Das Artareal umfaßt das Gebiet zwischen den Pässen von Portalé (im Westen) und Bonaigua (im Osten). Die Eidechse lebt in der alpinen Region, gewöhnlich in Höhen oberhalb 2000 m (zwischen 1700 m und 3062 m) und kommt dort in kleinen bis mittelgroßen, gewöhnlich eng begrenzten Populationen auf unterschiedlichem Gestein (Kalk, Schiefer, seltener und weniger dicht auf Gneis und Granit) und häufig in der Nähe von Seen und Bächen vor. Die Stichproben von Posets und Maladeta erscheinen als morphologisch zentral bzw. intermediär. An sie lassen sich die übrigen Stichproben aus verschiedenen Richtungen anschließen. Unter den gut repräsentierten Stichproben scheinen die von Bigorre und Ballibierna denen von Maladeta, Monte Perdido und Posets am nächsten zu stehen. Unter den weniger gut vertretenen Samples nähern sich die von Neouvielle und Arriel an Bigorre an. Punta Suelza Exemplare sind intermediär und nahezu nicht von M. Perdido und Posets Tieren unterscheidbar. Die Stichproben Maladeta, Besibierri, Aigüestortes and Ballibierna sind alle gleichermaßen gegenüber den intermediären Populationen differenziert. Insgesamt entsteht der Eindruck, daß die U-förmige Gebirgsformation von Posets, Maladeta und Ballibierna von einer Gruppe „zentraler Populationen“ bewohnt ist, von denen sich drei Stichproben morphologisch am stärksten abheben: Bigorre im Norden, M. Perdido westlich und Aigüestortes im Osten. Dieses Modell positioniert einen möglichen Refugialstandort der Art im Würm auf die Südhänge jenes U-förmigen Gebirgskomplexes, doch bleibt die Möglichkeit anderer kleiner Refugialräume, etwas im Gebiet von Aigüestortes. Das Szenario der Ausbreitung aus diesen Refugialräumen am Ende der letzten Vereisung wird dargestellt, eine das Gesamtmaterial einbeziehende vollständige Diagnose von I. bonnali wird gegeben.

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The study techniques permit us to check, from a morphological point of view, the existence of three well differentiated species in the Pyrenees: Iberolacerta aranica (Arribas, 1993), Iberolacerta aurelioi (Arribas, 1994) and Iberolacerta bonnali (Lantz 1927). ANOVA detects differences which accumulate aiming populations of different species. Characters with greater discriminative power among taxa are dorsalia, gularia, ventralia and the relative diameter of the tympanic scale. In the canonical analysis, first axis separates central-eastern pyrenaean populations (I. aurelioi) in respect to central pyrenaean ones (I. bonnali and I. aranica). Second axis separates central pyrenaean populations living in a small isolate at the northeast to the Garona river (I. aranica) from the western ones living in the main part of the pyrenean range (I. bonnali). I. aurelioi is characterized by high values ol ventral pattern, gularia, dorsalia and ventralia; and Iow values of masseteric and tympanic diameters, (temporal areas covered only by small tiny scales are not uncommon), generalyzed absence of contact among postocular and parietal scales and a lower number of circumanals and femoral pores. This species can be distinguished unequivocally also by his yellow belly, which is absent in the other pyrenaean species. I. aranica is characterized by greater diameters of the masseteric plate, relatively well developed ventral pattern. low values of dorsalia. ventralia, circumanalia and relatively short fore and hindlimbs. Usually shows a very characteristic scale disposition in temporal area, with three great scales (masseteric. tympanic and a great wedge-shaped intermediate scale). I. bonnali has intermediate pholidotic values among I. aurelioi and I. aranica. The three species show a good discrimination degree among them (superior to 75 %). The best discrimination is for I. aurelioi from which specimens rarely are mistaken with the other two species. Between I. aranica and I. bonnali there is a more imprecise discrimination, although ever superior to 75 % and more marked in males. In respect to the intraespecific variability among species from which more than one massif has been studied. I. aurelioi is the one which shows samples with the best discrimination degree among it (correct classification percentages from 71.42 to 91.6 %). In I. bonnali intracspecific discrimination is lower. Only the males from Bigorre reach the 75 % of correct classification, which suggests that this species is relatively uniform.

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Iberoalacerta aranica (ARRIBAS, 1993) ist ein Endemit der Zentral-Pyrenäen, der ausschließlich das Maubèrme Massiv und dessen Ausläufer zwischen Val d´Aràn (Spanien) und der Ariège (Frankreich) bewohnt. Vorkommen der Art sind in fünfundzwanzig 1 km x 1 km U.T.M. Rasterfeldern nachgewiesen; das mögliche Verbreitungsgebiet bedeckt maximal 36 derartige Rasterfelder. Die Eidechse bewohnt Schutthalden, Geröllfelder und steiniges Grasland (meist auf paläozoischen Schiefern und Kalken) im alpinen Vegetationsgürtel zwischen 1940 m (1900 m) und 2668 m Seehöhe. Alle untersuchten Populationen mit Ausnahme deren von Orlá scheinen untereinander mehr oder weniger in Kontakt zu stehen. In Abhängigkeit vom bewohnten Gesteinsuntergrund bestehen populationstypische Unterschiede in der Rückengrundfarbe. Tiere von Maubèrme ähneln jenen von Liat sehr; beide stehen den Serra d´Armeros Populationen (Barrados und Pica Palomera) nahe. Im allgemeinen dominieren in letzteren Populationen dunkle Farbtöne, während bei Exemplaren von Maubèrme und Liuat intermediäre oder hellere Grundfarben vorherrschen. Estany de Güerri ist ein Vorkommen am äußersten Ende der Serra d´Armeros, und das einzige von dort untersuchte Exemplar scheint morphologisch den Tieren von Armenos nahezustehen. Nur die Exemplare von Orlà unterscheiden sich deutlich von allen übrigen. Sie besitzen eine hellere Rückengrundfarbe mit deutlicher dunkler Färbung. Pholidotisch unterscheiden sie sich von den übrigen Populationen an der Körperunterseite (vor allem ihre größeren Gularia und Ventralia) sowie durch kleinere Masseterica und Analia; auch haben sie relativ längere Hinterbeine. Die Unterschiede in den Extremitätenproportionen könnten im Zusammenhang mit der gut ausgebildeten Kletterfähigkeit und dem bewohnten Gesteinstyp gesehen werden. Die Populationen von Orlà sind entweder tatsächlich isoliert oder sie stehen mit Maubèrme Tieren über einen sehr schmalen Korridor in Verbindung.

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Iberolacerta aurelioi (ARRIBAS, 1994) ist eine in ihrem Bestand gefährdete Eidechsenart mit einer sehr kleinräumigen Verbreitung. Wesentliche Merkmale ihrer Fortpflanzungsbiologie wie etwa die hohe Lebenserwartung (von bis zu 17 Jahren, die verzögerte Geschlechtsreife (nicht vor dem 5. Kalenderjahr bzw. 4 Überwinterungen) und sehr geringe Vermehrungsrate (ein Gelege aus 2 oder 3 Eiern pro Jahr) stehen im Zusammenhang mit der kurzen Jahresaktivitätsdauer in großen Höhen. Das Datum des Erscheinens nach der Winterruhe hing von den tatsächlichen Wetterverhältnissen im jeweiligen Jahr ab und war dann für alle Altersstufen und beide Geschlechter dasselbe. Die Paarung begann bald nach dem Verlassen der Winterquartiere (zweite Maihälfte bis erste Junihälfte). Für die Paarungsstellung war der Flankenbiß des Männchens charakteristisch. Die Eientwicklung erfolgte im Verlauf des Juni und die Eiablage fand zwischen Mitte Juni und Mitte Juli statt. Es erfolgte nur eine Eiablage pro Jahr, wobei (1) 2 oder 3 Eier gelegt wurden, welche Embryonen in fortgeschrittenen Reifestadien (Stadium 30 bis 32 nach DUFAURE et al. 1961) Bevorzugt wurden Ablageorte unter mittelgroßen bis großen flachen Steinen in leicht geneigtem Gelände. Unter solchen Steinen fanden sich zwei bis 12 frisch abglegte Eier, die von mehreren (im Durchschnitt zwei) Weibchen gemeinsam am selben Platz abgelegt worden sein mussten. Allerdings wurde solche Ablageorte häufig von Jahr zu Jahr gewechselt, sodaß nur wenige von ihnen in aufeinanderfolgenden Jahren benutzt waren. Die Veränderungen in den Abmessungen der Eier während der Inkubation werden beschrieben. Die Eizeitigung dauerte im Labor zwischen 31 bis 44 (im Mittel 35) Tage. Im Freiland traten die ersten Schlüpflinge zwischen Mitte August und Mitte September auf. Im Freiland zerstörte Fraß durch die Larve der Fliege Sarcophaga protuberans 25 % der Gelege. 64 % der im Labor erbrüteten Eier schlüpften. Der Schlupfvorgang und das frisch geschlüpfte Jungtier werden beschrieben.

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Iberolacerta bonnali (LANTZ, 1927 war von 07.00 bis etwa 16.00 Uhr GMT aktiv (gewöhnlich zwischen 09.00 und 11.00 Uhr), unabhängig von Geschlecht oder Alter. Die Orographie und das jeweils herrschende Wetter schränkten die nachmittägliche Aktivität der Eidechse ein. Im Laufe des Sommers und mit abnehmender Tageslänge verschob sich der Aktivitätsgipfel zunehmend in Richtung späterer Tagesstunden. In Kalksteingebieten begann und endete die Tagesaktivität früher als in Schiefergebieten, doch unterstützte in letzteren das tigmothermische Verhalten der Eidechsen wahrscheinlich den Prozeß des Aufwärmens. Die Hangneigung war nicht mit der Nutzungshäufigkeit durch I. bonnali korreliert. Die häufigsten und meist genutzten Mikrohabitate waren felsige oder grasbestandene Stellen. Die Eidechsen glichen Abweichungen von der mittleren Häufigkeit dieser Strukturen in allen drei Substrattypen (Kalk, Schiefer, Grandiorit/Granit) durch Bevorzugung oder Vermeidung der verfügbaren Mikrohabitate aus. In Schiefergebieten war Fels weniger häufig und wurde positiv selektioniert, während lose Gesteinbrocken relativ häufig waren und weniger angenommen wurden. Im Kalkgestein fehlte die Buschvegetation und die häufigsten Mikrohabitate, Fels und Gras, wurden etwas weniger angenommen, während umherliegende Steine selten auftraten und vermehrt als Aufenthaltsort gewählt wurden. Auf Grandiorit/Granit-Substrat wurden Grasstandorte weniger angenommen als Fels und loses Gestein. Die Körpertemperaturen aktiver Eidechsen reichten von 20.8°C bis 35.2°C (Mittel 28.96°C ± 0.24°C) und waren etwas stärker mit der Substrattemperatur (r = 0.37) als mit der Temperatur der Umgebungsluft korreliert (r = 0.33). Signifikante Unterschiede zwischen den Geschlechtern oder Altersgruppen oder im Hinblick auf unterschiedliche Substrattypen waren nicht feststellbar. Iberolacerta bonnali waren effektive Thermoregulatoren (die Anstiege der Regressionsgeraden von 0.14 bzw. 0.21 der Körpertemperatur in bezug zu Substrat- bzw. Lufttempetratur waren stark von I verschieden), aber nicht sehr präzise bei der Einstellung ihrer Körpertemperaturen (geringe Korrelationen) und damit eher Thermokonformisten (in einem breiten Temperaturspektrum aktiv), wie auch andere Iberolacerta-Arten.

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Der Autor legt eine vergleichende Untersuchung über Aktivität, Mikrohabitatwahl (Hangneigung, Flächenanteil von Fels, Steinen, nacktem Boden, Gebüsch und krautiger Vegetation im Lebensraum) und temperaturabhängiges Verhalten von Iberolacerta aranica (ARRIBAS, 1993), I. aurelioi (ARRIBAS; 1994) und I. bonnali (LANTZ, 1927) vor und beschreibt die Unterschiede in den untersuchten Größen im Hinblick auf Art, Geschlecht und Alter der Eidechsen. Im Aktivitätsmuster finden sich keine Unterschiede zwischen den Arten, doch differierten sie in mehreren Faktoren der Habitatausstattung ihres Aufenthaltsortes signifikant. Iberolacerta aurelioi wurde in steilerem Gelände als I. bonnali besonders aber als I. aranica festgestellt, doch waren die Unterschiede in der Hangneigung nich signifikant. Iberolacerta aurelioi und I. bonnali bevorzugten felsige Substrate und vermieden steinige Stellen (beide Geschlechter, alle Altersklassen) und nackte Böden (nur I. aurelioi). Iberolacerta aranica bevorzugte weniger felsigen als eher steinigen un d unbewachsenen Bodengrund. Büsche waren im Habitat von I. bonnali häufiger als in jenem von I. aurelioi und I. aranica. Krautige Pflanzen (Ganz allgemein alle Formen von Vegetation) fanden sich häufiger im Lebensraum von I. bonnali als an Stellen, wo I. aurelioi oder I. aranica vorkamen. Die drei Arten unterschieden sich voneinander in der Körpertemperatur (BT) und erwiesen sich als gute aber nicht sehr präzise Thermoregulatoren (besonders gegenüber den Subastrattemperaturen) sowie mäßige Thermokonformisten. Die Körpertemperatur war bei Iberolacerta aurelioi niedriger als bei I. bonnali (p < 0.05) und besonders I. aranica (p < 0.01). Die mittlere Körpertemperatur betrug bei I. bonnali 29.20 ± 0.30°C (22.8 – 35.2°C; n = 64), bei I. aranica 29.49 ± 0.20°C (22 – 36.5°C; n = 143) und bei I. aurelioi 28.13 ± 0.32°C (16.6 – 34.4°C; n = 109). Die Erwärmung schien bei I. aurelioi und I. bonnali hauptsächlich heliothermisch, bei I. aranica tigmothermisch zu erfolgen.

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Canonical Discriminant (CDA), ANOVA and ANOSIM analyses were calculated for all recen tly known distribution areas of Iberolacerta cyreni including severtal small and unstudied peripheral populations. The monly differentiated sample is Guadarrama (the nominate subspecies), with very limited overlap in the CDA (correct classification > 70%) and different from nearly all the other samples in ANOSIM. Guadarrama is a recently differentiated but well diagnosable (morpho) subspecies (with lower values of dorsalia, ventralia and greater values of circumanalia). Despite the mtDNA differences of the Béjar specimens, their morphology is largely equivalent to that of I. Cyreni castiliana (Gredos), but clearly differ in their female body elongation (near 1 cm) with shorter limnbs, a possible strategy to increase clutch size. Populations from the Sierras de Avila (Villafranca, Serrota and Paramera) are very similar among them. Villafranca (in males) together with Béjar (in females) are the most connected samples in MST, and the root of the species differentiation from a morphological point of view, once discarded geographical and climatic influence on morphology. All populations except Guadarrama shall be considered as I. c. castiliana by their morphological identity with gredos. These morphological similarities probably are the reflect of extensive gene flow among them, responsible of maintaining their morphology largely euqivalent.

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In this paper we deal on the ultraviolet color (invisible to us): where we can find it, the capability of animals to see it and the advantages that this color perception offers to them. As the simplest way to detect it is the photography, we describe and review how to photograph the UV, as a result of 15 years of amateur experience, searching and testing nearly in complete blindness due to the lack of practical information about “how to do it”. We describe the different kinds of photography (chemical and digital); the cameras and objectives suitable (both astronomically expensive ones and cheap options); what are the best characteristics that the objectives should have for this purpose; the films suitable for their use in chemical photography; the different filters (current or discontinued) manufactured along the years; and the subtle combinations among the different materials to obtain pure UV photographs. This kind of scientific photography is mainly used in forensics, forgery detection, art dermatology and less in Natural History, despite the fact that a great part of animals see this color and use it in important questions of their biology as the social behavior, mate choice or the food search.

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Zur Naturgeschichte von Iberolacerta martinezricai (ARRIBAS, 1996), der seltensten Reptilienart Kointinentaleuropas, liegen nur wenige Angaben vor. Das zahlenmäßige Geschlechterverhältnis betrag nach Informationen aus der Datenbank des Autors 0,74:1 (Männchen: Weibchen), während im deutlich größeren datensatz der feldbeobachtungen (dieser schließt allerdings Beobachtungswiederholungen ein) die Männchen überwogen (1,27:1). Alla Altersgruppen und Geschlechter zeigten ähnliche, die Morgenstunden betonende Tagsaktivitätszyklen. Nachmittagsaktivität erfolgte gehäuft an schattingen Orten, wobei Sonnenexposition bei hohen Lufttemperaturen vermiden wurde. Zwei Häutungsperioden (eine im April, eine um Ende Jun und Juli) wrden beobachtet. Der Lebensraum war mäßig neiegtes, ausnahmslos von Felstrümmern und großen Blöcken, mit reichlich Moos- und Flechtenbewuchs dominiertes Gelände, was auf erhöhte Feuchte und geologische Hang-Stabilität hinwies. Besondere Vorlieben fürf einzelne Mikrohabitate waren nicht feststellbar. Die Körpertemperatur aktiver Individuen war die höchste innerhalb der Gattung Iberolacerta: 33.31±0.12°C (Spannweite 22.8-39.2°C) ohne signifikanten Geschlechtsunterschied. Im Mittel hatten blaufärbige Männchen eine höhere Körpertemperatur als grüne (33.73±0.19°C gegenüber 32.8±0.28°C). Die Aufenthaltsorte adulter Weibchen wiesen eine höhere Lufttemperatur auf (23.77±0.20°C, Spannweite 13.8-37.1°C) als die der Jungtiere. Dioe Substrattemperaturen (30.08±0.31°C, Spannweite 16.5-52.1°C) waren an den Aufenthaltsorten derWeibchen höhetr als an dene der Männchen. Iberolacerta martinezricai war mäßig thermokonformistisch (Spannweite der Aktivitätstemperatur: 16.4°C) und ein effektiver und ziemlich präziser Thermoregulator was die Substrattemperatur betrifft(r = 0.36, Anstieg 0.14), aber zeigte kein thermoregulatorisches Verhalten hinsichtlich der Lufttemperatur (r = 0.15, Anstieg = 0.09). Tigmothermie, Aktivität im Schatten bei hoher Lufttemperatur und Aestivation werden diskutiert-

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Sex dimorphism: Males and females of Iberolacerta martinezricai differ at p < 0.001 in Ventralia (MF), Forelimbs length (M>F), Hindlimb length (M>F), Pileus length (M>F) and at p<0.05 in Collaria (M>F) and Dorsalia (M>F). Batuecan lizards are overall less di-morphic than the three species of Pyrenean rock lizards. Morphological variability: Variability Index calculated (ΣCV) for I. martinezricai was 6.8 in males and 6.0 in females. Results are very similar (although slightly smaller) to the obtained in the Pyrenean Iberolacerta spp. From two chosen characters, variation coefficient in Gularia falls also between the normal values for the bisexual species and it is a bit greater than in parthenogenetic species; or it is clearly greater as in Dorsalia. Nothing suggests a drastic reduction of the variability in the Batuecan lizard but some characters recall for a relatively small morphologic variability in some characters, in parallel with the lack of mitochondrial variability, perhaps due to its reduced effectives. Allometry: Both in males and females, the most representative character of growth is the body length (SVL). Most other characters match very closely to this SVL with very similar coefficients: limbs, Parietal, Pileus length and the Anal plate width. Masseteric, Tympanic and Anal length also grow, but not in parallel with SVL. Anal length grows considerably and independently of body size. In the case of fe-males, body limbs have a negative allometry respect to female’s body length and Masseteric, and especially Tympanic plates have a negative allometry respect to the total growth. Parallel variation in escalation: In males the most related characters of escalation are the two rows of Femoralia, and two characters that have contrary signs: Supraciliar Granula (but not very marked, as coefficients are very small) and specially Circumanalia (that varies with total independence). Also there is a small relationship of Supraciliar Granula with Gularia and even more distant with 4th-toe Lamellae and Circumanalia. In females on one side (first axis), both Supraciliar Granula rows are related to Femoral pores, 4th toe Lamellae, Ventralia and Femoralia; and on the other side (second axis) both sided Femoralia are lonely related with Dorsalia, Collaria and Gularia, and just inversely related to Supraciliar Granula. Sexual maturation is probably reached when lizards are three years old (in his 4CY). Our smallest gravid female was 56.7 mm, and was probably reproducing in the spring of their 5 CY (when 4 years old). Al-though it is possible that some specimens (especially males, but also some females) reach maturity during 3CY, as it occurs in I. monticola. Growth rate (K) in the first years is 0.145696 ± 0.0633135 (95% confidence). Calculated length at infinite (maximum size if the same rate or growth curve was ap-plied throughout the lifetime of the animal) would be: L∞ = 70.036 ± 13.7598 (95%). This estimate of the asymptotic size (70.036 mm) is very close to the true maximum size found in the total sample of adults (69.31 mm). Growth decreases progressively and the resulting curve is fairly straight, being the growth not far from linear and very similar in both sexes. Longevity: The biggest male (SVL 68.15 mm) and the biggest female conserved (SVL 68.86 mm) had 7 growth rings (two big and five narrow) in the femur. The two wide rings correspond to the two first years of rapid growth (2CY and 3CY), but when lizards reach sex maturation growth decreases, being latter rings very narrow and even difficult to count. Both specimens had presumably 8 years old (they are 9CY animals: the birth year and 7 growths seasons, and captured just in the beginning of their 9 CY). Tail autotomy: Iberolacerta martinezricai hatchlings are able to break their tails from birth. Observed hatchlings (1CY) in nature had intact tails. In 2CY, 3CY and 4CY already appear broken and regenerated tails in some specimens. In adults, there are less than a quarter of specimens with apparently unbroken tails, and the others fall in the different categories of regenerated ones (tail tip, middle of the tail or the whole tail from basis) in proportions similar to one quarter. There are no differences between both sexes in the different categories of broken tails, but there are significant differences between juveniles and adults. Females have a slightly big percentage of broken tails from its basis. Bright colored tail in juveniles (bluish in I. martinezricai), is lost during their 2CY. It seems to be an antipredatory strategy. Hatchlings of the Batuecan lizard are able to attract attention on its tail by a striking whirl of its distal part, unlike adults that are unable with their stiffer tail (which can move only in very limited cases of danger or excitement).

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Tail regeneration in 706 Iberolacerta (Pyrenesaura): 300 I. bonnali, 212 I. aranica and 194 I. aurelioi was studied, distinguishing four situations: 0) intact tail; 1) distal part (point) of the tail regenerated; 2) Tail regenerated from its half part and 3) tail broken and regenerated from its basis. The incidence of autotomy (tail break) in Iberolacerta (Pyrenesaura) runs in adults from a 47.5% up to a 68.3%. In juveniles is sensibly lower and goes from 10.9% to 23.8%.In I. bonnali percentage of autotomized tails is almost identical in both sexes (48% in males and 47.5% in females), whereas in juveniles is only 22.4%, without differences between adults but yes between adults and juveniles. In I. aranica, percentage of autotomized tails is 68.3% in males (fairly high), a 55.3% in females and a 10.9% in immatures. There are not significant differences between males and females (despite to be near to significance) but exist between adults and juveniles. In I. aurelioi males have a break percentage of 54.4%, females 59.8% and juveniles 23.8%, without differences between males and females, and contrary to the other two Pyrenean species, also inexistent between adults and juveniles. Comparing the three species among them, only males differ (the difference arises between I. bonnali and I. aranica), and all the other comparisons among females and juveniles are not significant. Juveniles of I. aurelioi (only any population of I. aranica or rarely and few vivid in I. bonnali) present bluish of greenish tails that do not broke until some days after birth and are able to wave sinuously in order to attract the predators attention upon it. Also, the known and possible predators are reviewed.

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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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Der Autor untersuchte das Fortpflanzungsgeschehen von Iberolacerta martinezricai (ARRIBAS, 1996). Kopulationen begannen Ende April. Im Juli trugen alle Weibchen zahlreiche frische Paarungsnarben auf Bauch und Schwanzbasis. Iberolacerta martinezricai ist monoestrisch. Die Eiablage erfolge im Juli und Anfang August, am häufigsten gegen Ende Juli. Das Eiablageverhalten wird beschrieben. Eier wurden zwischen 9:00 und 15.30 Uhr (GMT) gelegt, wobei der ganze Ablagevorgang 33-42 Minuten (Durchschnitt: 36:57 Minuten) dauerte. Die Kopf-Rumpf-Länge reproduktiver Weibchen betrug 56.7 – 69.71 mm. Die Gelege umfaßten zwei bis sechs Eier (Median und Modalwert: 4, Mittel: 4,23). Die abgelegten Eier klebten nicht aneinander. Biometrische Daten von 49 Eiern (darunter 12 Laboraufzuchten) werden angegeben. Während des Großteils der Inkubationsphase nahmen Eivolumen und Eimasse durch Wasseraufnahme kontinuierlich zu. Zu Begin der Inkubation erfolgte ein rascher Anstieg des Ei-Querdurchmessers (dieser wurde während der Ablage durch die Enge des Geburtskanals verringert). Die Inkubation dauerte 33-42Tage (im Mittel 37.75 Tage). Das Schlüpfen erfolgte zwischen 3:00 und 22:00 Uhr (GMT) (im Mittel 9.40 Uhr) und dauerte 4.10 – 12 Stunden (im Mittel 6.9 Stunden). Die Merkmale des Schlüpflings werden beschrieben. Der Eizahn bestand für acht bis 36 Stunden (im Mittel 23.6 Stunden nach Schlupfbeginn. Schlüpflinge können von Geburt an mit ihrem leuchtend blauen Schwanz wedeln und diesen autotomieren. Diese auffällige Schwanzfärbung verschwand im Laufe des zweiten Kalender-Lebensjahres.

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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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Busack et al. (2016) concluded that the name Darevskia should be attributed to Arribas (1999) instead of Arribas (1997), rendering the latter name a junior synonym of Caucasilacerta Harris et al. (1998). In this paper, we provide a detailed answer to Busack et al. (2016) and argue that Arribas (1997) fulfilled the publication requirements of the International Code of Zoological Nomenclature (ICZN) in place in 1997 and, therefore, Darevskia Arribas, 1997 is a valid name. In order to clarify this issue, a case has been submitted to the ICZN. To avoid unnecessary confusion in Old World Lacertid taxonomy, we recommend that until the commission rules on this case, Arribas (1997) should continue to be considered the authority of both Darevskia and Iberolacerta.

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Iberolacerta cyreni martinezricai is elevated to the species level (I. martinezricai) based on both morphological and molecular data. The phylogenetic analysis using two mitochondrial and one nuclear gene shows I. martinezricai is more closely related to I. monticola than to I. cyreni. A multivariate analysis of the morphological data also supports the affinities between I. martinezricai and I. monticola but, at the same time, clearly indicates that I. martinezricai is morphologically distinct from both I. monticola and I. cyreni. The molecular data suggests I. cyreni and the clade formed by I. monticola + I. martinezricai split approximately 6.1 Mya, during the Mesinian Salinity Crisis, when climatic conditions around the Mediterranean area changed dramatically as a result of the desiccation of the Mediterranean Sea. Separation between I. martinezricai and I. monticola occurred approximately 2 Mya but, with at least two equally plausible alternative hypotheses, their biogeography is still unclear. New data on the habitat and distribution of I. martinezricai indicates its distribution area is very small (12–15 km2), and that it lives in a climatically extreme habitat for this kind of mountain species. As a result of that and the low numbers of individuals, I. martinezricai is considered here as Critically Endangered.

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A new species of Iberolacerta is described from the Montes de León (northwest Iberia). This new species, Iberolacerta galani sp. nov., is characterized by its relatively large size, high number of blue ocelli on the shoulders and the relatively frequent contact or near-contact between the supranasal and the first loreal scale, the fairly straight squamosal bone (only curved on its posterior part), a unique karyotype in Iberolacerta combining 2n=36 chromosomes, an L-type NOR and differentiated W and Z sex chromosomes, and unique mitochondrial DNA sequences for the cytochrome b and 12S rRNA genes. The correlation analyses show that morphology in general, but especially scalation, is strongly correlated with the amount of precipitation during the months of lizard activity, which suggests that these are not good taxonomic characters, and that other characters apparently independent of the climate like for instance osteological, karyological and DNA features are much more reliable in delimiting species boundaries in Iberolacerta. According to our phylogenetic analyses, I. galani nov. is part of a very well supported clade that originated around 2.5 mya and also includes I. monticola and I. martinezricai. Phylogeny suggests I.martinezricai might be the sister taxon to I. galani nov. from which it split approximately 2 mya, at the beginning of the Pleistocene. The clade containing I. galani nov., I. martinezricai and I. monticola was probably widely distributed across western Iberia during moderately cool and moist phases of the Pleistocene, but it was probably restricted to its present range as a result of the general temperature increase during the Holocene and competition with other lacertid lizards. Iberolacerta galani nov. is endemic to the Montes de León, where it is isolated from the other species of the “monticola-group” by the Duero and Miño-Sil Rivers, but particularly by the Bibei river valley.

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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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The three lacertid lizards species of the Iberolacerta genus, which have recently been described or recognised as different species, are reptiles that live strictly at the highest altitudes in Europe, from elevations of near 1900 up to more than 3000 m a.s.l. in the Pyrenees (Spain, Andorra, France). In this paper, the reproductive cycle and reproductive characteristics of these species are described for the first time. Data were obtained from field studies carried out in different high-mountain Pyrenean locations from 1989–2002. In addition we also conducted hatching studies in laboratory. Due to the harsh climatic conditions in the high mountains the activity cycle of these species is very short, just over 4 months, from mid-May to late September or beginning of October. This affects their reproductive cycle (i.e., only one annual egg-clutch is produced), as well as other reproductive characteristics, as the existence of a very advanced embryonic development at oviposition, which is interpreted as an advanced stage in the tendency towards viviparity, and diverse life history characteristics as a very little annual growth which greatly delays sexual maturity to 4 years in males and 4–5 years in females. Clutch size correlates significantly with female snout-vent length (SVL) in all three species. The average clutch size is 2.53 eggs in I. aurelioi (the smallest species), 3.03 in I. bonnali and 3.44 in I. aranica (the biggest species). Notwithstanding the differences in egg number the three species have a similar egg volume. The incubation period in the laboratory is very short with an average of 30–36 days among the species. The low reproductive potential observed in these three species and revealed in this paper is a strongly threatening factor which, together with their extremely reduced distribution area, endangers these endemic, rare and very threatened species.

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Iberolacerta (cyreni) martinezricai has a karyotype of 2n = 36, all acrocentric macrochromosomes. Nucleolar Organizer Regions (NORs) are interstitially situated on a medium small (MS-type) chromosome, a derived character shared with I. cyreni, which differs in details of W-sex chromosome: W is heteromorphic and heterochromatic in I. cyreni, while it is homomorphic and euchromatic in I. (c.) martinezricai. Osteologically, it is characterized by the presence of a triangular-shaped expansion of the squamosal towards the supratemporal fenestra, and by the presence of seven (instead of six) posterior short ribbed presacral vertebrae. These odd characters probably became fixed in this relic taxon by imbreeding. Its differences and affinities are discussed, proposing a specific status for this taxon.

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Kroatien ist eines der herpetologisch artenreichsten Länder Europas und daher ein beliebtes Reiseziel mitteleuropäischer Herpetologen. Aufgrund des in den letzten Jahrzehnten stetig weiter ausgebauten Autobahnnetzes ist das Land gut und schnell zu bereisen, lediglich die Grenzwartezeiten an den Urlaubswochenenden sind noch einigermaßen mühsam, da Kroatien noch nicht im Schengen-Raum aufgenommen wurde. Viele im Rahmen der ÖGH oder Herpetofauna.at durchgeführten Exkursionen führten uns in den letzten Jahren nach Kroatien.

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Animals communicate via a variety of sensory channels and signals. Studies on acoustic and visual communication systems suggest that differences in the physical environment contribute to the variety of signalling behaviour, with species investing in those signals that are transmitted best under the local conditions. Whether or not environmental tuning also occurs in chemical communication systems has received much less attention. In the present study, we examined the effect of several aspects of the physical environment on the chemical communication system of lacertid lizards (family Lacertidae). The numbers of femoral pores are used as a proxy reflecting how much a particular species invests in and relies upon chemical signalling. Femoral pores are specialized epidermal structures that function as a secretion channel for the waxy substance produced by glands. In some lacertid species, the secretion carries infochemicals that play an important role in social communication. The number of femoral pores varies considerably among species. We have compiled data on femoral pore numbers for 162 species and tested for the effects of climate and substrate use. After correcting for body size and taking the phylogenetic relationships among the species into account, we found no effect of climate conditions or latitude on species pore numbers. Substrate use did affect pore numbers: shrub-climbing species tended to have fewer femoral pores than species inhabiting other substrates.

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Lizards communicate with others via chemical signals, the composition of which may vary among species. Although the selective pressures and constraints affecting chemical signal diversity at the species level remain poorly understood, the possible role of diet has been largely neglected. The chemical signals of many lizards originate from the femoral glands that exude a mixture of semiochemicals, and may be used in a variety of contexts. We analyzed the lipophilic fraction of the glandular secretions of 45 species of lacertid lizard species by gas chromatography/mass spectrometry. The proportions of nine major chemical classes (alcohols, aldehydes, fatty acids, furanones, ketones, steroids, terpenoids, tocopherols and waxy esters), the relative contributions of these different classes (‘chemical diversity’), and the total number of different lipophilic compounds (‘chemical richness’) varied greatly among species.We examined whether interspecific differences in these chemical variables could be coupled to interspecific variation in diet using data from the literature. In addition, we compared chemical signal composition among species that almost never, occasionally, or often eat plant material. We found little support for the hypothesis that the chemical profile of a given species’ secretion depends on the type of food consumed. Diet breadth did not correlate with chemical diversity or richness. The amount of plants or ants consumed did not affect the relative contribution of any of the nine major chemical classes to the secretion. Chemical diversity did not differ among lizards with different levels of plant consumption; however, chemical richness was low in species with an exclusive arthropod diet, suggesting that incorporating plants in the diet enables lizards to increase the number of compounds allocated to secretions, likely because a (partly) herbivorous diet allows them to include compounds of plant origin that are unavailable in animal prey. Still, overall, diet appears a relatively poor predictor of interspecific differences in the broad chemical profiles of secretions of lacertid lizards.

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The authors give a description of the herpetofauna of some area´s in Yougoslavia they visited in june 1979. Some attention is given to the habitat of several species. The differences between certain island and mainland populations of some Lacerta- and Podarcis-species are discussed to some extend.

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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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In connectivity conservation and ecological network planning, the selection of focal fragmentationsensitive species represents a priority step. Nevertheless, despite their strategic role, selection of focal species has traditionally been carried out using charismatic and/or non objective approaches. In this way, actions of planning and conservation could be ineffective. Using as a case study Italian reptiles, we apply an expert-based approach for the selection of focal species on the basis of sensitivity to components of habitat fragmentation (habitat area reduction, increase of habitat isolation, increase of edge effect and landscape matrix disturbance) and of intrinsic ecological traits of the species (trophic level, dispersal ability, body size, niche breadth, rarity). The threshold values for each component of fragmentation defined a set of 21 focal species that can be divided into the three macro-components of human-induced habitat fragmentation (HIHF) towards which they show a sensitivity, the suitable spatial scale of populations and relative suitable habitat categories. Among these species, seven can be sampled easily with standard, low-cost field protocols. The selected species largely coincide with the species known in literature as fragmentation-sensitive.

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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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Understanding the evolutionary background of consistent individual differences in behaviour (animal personality in single, behavioural syndrome across multiple behaviours) is a prime aim of current evolutionary behavioural ecology. Properly assessing true fitness is notoriously hard, especially in the wild. One way to circumvent this problem is to use quality-indicator traits as fitness proxies. Hence, studying the link between behavioural consistency and quality-indicator traits can be an important step in the evolutionary analysis of behavioural consistency. In the study outlined in my thesis, we studied whether wild-caught, adult male Iberolacerta cyrenii (Squamata: Lacertidae) had personality and showed behavioural syndrome, and whether behavioural consistency was linked to potentially quality-indicator traits. We analysed both behavioural types (mean behaviour) and individual behavioural variation (standard deviation around the mean). Lizards in the study population showed high behavioural consistency both within and across activity and risk-taking, indicating the presence of animal personality and behavioural syndrome. More active individuals were less consistent than their passive conspecifics, while more risk- taking lizards were more consistent than the risk-averse ones. Activity was positively related to total brightness, while risk-taking was positively related to relative tibia length and number of femoral pores. The complex behavioural variable including both activity and risk-taking was positively related to relative tibia length and number of femoral pores, but negatively to the number of lateral blue spots. Consistency of risk-taking showed positive condition-dependence. Our results are pioneering in reptiles, which are a promising model for the future. Due to the correlative nature of our study, making clear cause-causative interpretations of the numerous behaviour – potential quality-indicator trait links is impossible, they are still suggestive of important evolutionary mechanism. Hence, our results form a springboard for targeted manipulative experiments in the future.

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The recognition that consistent between-individual behavioural differences (i.e. animal personality) are biologically relevant and valid phenomena arrived only in the last ca. 15 years to the field of behavioural ecology. The existence of animal personality is not intuitively expected, since it can decrease fitness in several ways. For instance, aggressive individuals stay aggressive in contexts or situations when this behaviour is highly maladaptive, like against their mate or offspring, or in the presence of predators. Thus, the identification of mechanisms playing role in the development and evolution of animal personality has an utmost importance. Ecological and physiological factors are known to have the potential to create long-term between-individual behavioural differences, especially during the early stages of ontogeny. However, the role of these factors during adulthood is rather controversial. Carpetane rock lizard (Iberolacerta cyreni) is a small diurnal lacertid, endemic to the high-mountainous regions of Central Spain, and one of the best studied lizards in Europe. Using this species as a model organism I aimed to study how testosterone treatment (testosterone vs. placebo) affects aggression of wild-caught reproductive adult male I. cyreni in two ecologically distinct situations (perceived predation risk present vs. absent). I focussed on two components of behavioural variation: individual mean behaviour and the strength of behavioural consistency. The applied treatments affected only behavioural consistency: control males (placebo treatment × predator absent) lacked aggressive personality, i.e. individuals were statistically indistinguishable, while testosterone treatment and/or the presence of perceived predation risk resulted in strong personalities. My results suggest that consistent individual behavioural variation is not necessary stable and relevant ecological or physiological stimuli are necessary for the emergence of animal personality.

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Eight polymorphic microsatellite loci are described for the Iberian rock lizard species, Iberolacerta cyreni. Loci were isolated from a partial genomic library that had been enriched for AAAG repeat sequence. Number of alleles per locus ranged from two to 13 in a sample of 24 individuals from the Sierra de Guadarrama (central Spain). Observed heterozygosity ranged from 0.13 to 0.96. At least three loci were amplified and polymorphic in four other Iberian rock lizard species: Iberolacerta monticola, Iberolacerta bonnali, Iberolacerta aranica and Iberolacerta aurelioi. These markers will be used to study mating strategies and reproductive success in I. cyreni.

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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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The Herpetological Collection of the Trieste Natural History Museum has almost 700 specimens of European Sauria belonging to about 33 taxa, including all the lacertids of northern Adriatic re- gions. The origin of the collection dates back to the foundation of the Museum of Trieste in 1846, but it was increased mainly between 1871 and 1939. The Herpetological Collection of the Trieste Natural History Museum has a great scientific and historical importance; it documents the varia- tion of the composition and distribution of northern Adriatic Sauria fauna during this century, in- cluding endemic varieties typical of little islands.

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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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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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A doctoral thesis presented in the late 1990s by óscar J. Arribas has been frequently cited as the authority for the nomenclatural status of two generic names in the lizard family Lacertidae, Darevskia and Iberolacerta. These names were again introduced, along with the addition of the subgeneric name Pyrenesaura, in 1999 via publication in an international, peer-reviewed journal. We discuss nomenclatural aspects of how these taxon names were presented, detail inconsistencies in the use of publication dates associated with these names, and resolve nomenclatural problems by demonstrating that the date of their first publication, in compliance with the International Code of Zoological Nomenclature, was 1999. This renders the name Darevskia Arribas, 1999 a junior invalid synonym of Caucasilacerta Harris, Arnold & Thomas, 1998, which, contrary to previous assessment, is not a nomen nudum, and must replace Darevskia as the valid name for this genus.

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Im Untersuchungsgebiet in den Lienzer Dolomiten (Kärnten, Österreich) kommen Iberolacerta horvathi (MÉHELY, 1904) und Podarcis muralis (LAURENTI, 1768) zwischen 714 und 1285 m ü. M. sympatrisch, zwischen 919 und 1057 m syntop vor. Rund 10 % der Eidechsenstandorte sind von beiden Arten etwa 50 % bzw. 40 % nur von P. muralis bzw. I. horvathi besetzt. Die beobachteten Individuendichten liegen bei I. horvathi mehrfach höher als bei P. muralis. Die felsig-steinigen Eidechsenhabitate – vom glatten, steilen Fels bis zur porösen, flachen Schutthalde – wurden nach ihrer Kompaktheit, Höhe, Neigung, Spaltenbildung und Oberflächenstruktur vier Typen zugeordnet, zu deren Besiedlung unterschiedliche Kletterfähigkeiten erforderlich sind. Auf dem glatten steilen Felsformationen kam nur I. horvathi vor, auf Schuttflächen nahezu ausschließlich P. muralis, Übergangsformen konnten beide Arten beherbergen.

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

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

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The dietary habits of Lacerta horvathi MÉHELY, 1904, from the Tarvisio Forest (Carnic Alps, Province of Udine) were studied using faecal pellet analysis. 81 specimens were analyzed for prey remnants in tlieir faecal pellets. The sexes did not differ significantly in terms of SVL (one-way ANOVA: P < 0.30), but males were larger than females. Lacerta horvathi fed mainly upon Opiliones and Araneae, while the proportion of insects was quite low. Both mean number of prey items per faecal pellet and food niche breadth was higher in males than in females and juveniles, but the differences between the three classes were not statistically significant. Compared with several other lacertid species, L. horvathi seems to be much less a food generalist. In fact, it is strongly specialized in preying upon grounddwelling arthropods. Furthermore, the prey types eaten suggest that this lizard is an active forager, as expected in small-sized rupicolous lacertids.

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Flight initiation distance in relation to substratum type, sex, reproductive status and tail condition was studied in two lacertid lizards with contrasting habits: the ground-dwelling common lizard Zootoca vivipara and the rupicolous Horvath`s rock lizard Iberolacerta horvathi. These species were studied in sympatric populations in a mountain area in North-Eastern Italy, Tarvisio Forest. Mean escape distance was significantly higher in I. horvathi than in Z. vivipara. In both species there were significant differences between sexes, with males escaping at longer distances than females but there were no significant differences between adults and subadults. In both species there were no differences in escape distance of females in different reproductive states. In Z. vivipara specimens with broken tails escaped at a shorter distance than individuals with intact tails. Substratum type had a significant effect on escape distance in both species.

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The chromosomes of Lacerta horvfthi have been studied by means of conventional, C-banding, and silver-NOR techniques. The karyotype of this species, characterized by 36 acrocentric macrochromo- somes, lacks the typical pair of microchromosomes shared by all other lacertid lizards. It is hypothesized that the microchromosomes could have been translocated to the large elements of the karyotype. The occurrence of such a rearrangement in the chromosome complement ofL. horvhthi underlines its isolation from the other species of the subgenus Archaeolacerta. The C-banding analysis evidences the existence of a female sex heteromorphism in which the W-chromosome has the same shape and size of the Z, but differs from it in being completely heterochromatic. The nucleolar organizer regions (NORs) are located on a pair of medium size chromosomes in subtelomeric position, where the standard Giemsa-staining reveals secondary constrictions.

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seven species of the rock lizard genus Iberolacerta are represented on the iberian Peninsula. These lizards are considered to be under threat of extinction, with Iberolacerta martinezricai (ArribAs, 1996), being one of the most endangered reptiles in Europe. There is however, lack of knowledge about its conservation status, distribu- tion and ecology. The authors’ surveys in 2007 and 2008 aimed at clarifying the distribution status of the species in Central spain, where 63 uTM squares (1 km x 1 km in size) were sampled and the density of the lizards, their distribution and habitat preferences studied. Iberolacerta martinezricai was detected in 23 out of the 63 uTM squares (36.5 %), with densities ranging from 25 to 50 lizards/ha. The statistical analysis revealed that the presence of the rock lizards was associated with altitude, lichen cover and rock/boulder size. Accordingly, the species is restricted to rocky slopes of the Peña de Francia mountain range. The results clearly indicate that the range of the species is highly restricted, that the habitats occupied are very specific, and that the population size is low compared with other species of the genus Iberolacerta. based on these findings, the species is categorized as Critically Endangered (Cr) under the criteria of the international union for the Conservation of nature (iuCn).

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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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The Horvath’s rock lizard, Iberolacerta horvathi, is occurring from 200 to 2000 m a.s.l. on the eastern Alps (NE Italy, S Austria, NW Slovenia) and the northern Dinaric Mountains (central and S Slovenia and NW Croatia). This species is the only representative of Iberolacerta outside Iberian Peninsula and currently shows a discontinuous occupational range. We collected tissue samples and extracted total genomic DNA of 110 samples from 23 localities across its entire distributional range. We grouped localities into three main geographical areas: northwestern (NW), central and southeastern (SE). For all samples we amplified a fragment of the mtDNA cytochrome b (Cytb) gene whereas a fragment of the nuclear melanocortin 1 receptor (MC1R) gene was amplified for a subset of 53 samples. Despite the relatively narrow geographical scale, the Cytb haplotype network indicated a high number of unique haplotypes present in each area (6 in the NW and 14 in the central and SE). In NW and in central area we detected two shared haplotypes: a) the widely distributed H5, and b) the H21, shared by only two populations. On the contrary, SE populations did not share any Cytb haplotype with other areas. The nuclear haplotype network analysis gave similar results, where multiple private alleles occurred in each area (4 in NW and central, and 3 in SE) and a widespread haplotype shared across the three areas. Results suggest that current distributional fragmentation is partially mirrored by the phylogeographic history of the species, with private alleles occurring in each of the main geographic areas, indicating limited gene flow among populations at least in the recent past. Knowledge of genetic structure of this endemic species provides essential information to orient activities to ensure its future conservation.

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The Horvath’s rock lizard Iberolacerta horvathi (Méhely, 1904) is an understudied lacertid species, which is geographically isolated from its congeners and currently classified as Near Threatened under IUCN red listing criteria. Due to its limited distribution and narrow environmental preferences, we expect that the species had been affected by past climatic oscillations. By combining all available data on species occurrences and our own records, we produced an updated distribution map. We used ecological niche models to identify the current environmental factors underlying the species range and developed a habitat suitability map. We sequenced one mitochondrial and one nuclear marker to characterize the distribution of the genetic variability and infer the historical demography of the species. The habitat suitability map identified areas where targeted field searches should be prioritized, as well as unsuitable habitats coinciding with likely barriers to gene flow. We found considerable genetic variability suggesting that the species probably survived the Pleistocene glaciations in at least two main refugia, one in the South and the other in the central/northern portion of its distribution. While southern populations show a moderate demographic decrease starting at the onset of the Eemian interglacial, the northern populations underwent an expansion during Late Pleistocene. We provide a revised species distribution and a first characterization of its genetic variability across its distribution to guide conservation priorities for this endemic and spatially restricted species.

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When a predator is immobile near an immobile prey, the probability that the predator will detect and attack increases over time. The prey’s cost of moving, thereby abandoning crypsis due to immobility, therefore decreases. Cost of not moving increases over time if movement is required for prey to conduct fitness-enhancing activities. We tested a cost-benefit model that predicts effects of factors that affect predation risk and cost of not moving on a prey’s latency to flee. Acting as simulated predators, we conducted experiments on the lizards Iberolacerta cyreni and Podarcis lilfordi. All predictions for 5 risk factors and a cost of moving factor were verified. Lizards fled sooner when the predator stood closer, approached rapidly rather than slowly before stopping, approached directly rather than indirectly, and gazed at the lizard rather than away from it, and after the second of 2 successive approaches. Latency to move was shorter in the presence than absence of a mealworm, suggesting the importance of opportunity cost of immobility. The effect of standing distance has 2 components, greater rate of detection by the predator and greater risk of being captured if detected at shorter distances. Escape theory has been highly successful in predicting how close a prey allows a predator to approach before fleeing. Our model extends an economic approach to study of escape decisions in response to an immobile predator that may be an ambush forager or an active forager that has stopped moving nearby.

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

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Our thesis aimedto recognize which intraerythrocytic blood parasites infect two species of Slovene lacertids, common wall lizard (Podarcis muralis),and Horvath`s rock lizard (Iberolacerta horvathi). We confirmed the identity of the parasites using molecular approaches by sequencing a part of the 18S rRNA gene. All observedparasiteindividuals belong to the genus Karyolysus, knownby parasitizing reptiles and lacertids specifically.Karyolysus is a genus of protozoan hamogreagrines.Further work included the analysis of prevalence and intensity of parasitic infection in both species of lizards. We compared prevalence and intensity between species, sexes, size, locations, altitudes,and whether the populations live in syntopy or alotopy. We confirmed that the prevalence and intensity of blood parasites were significantly higher in Horvath`s rock lizards. We also observed a higher prevalence of parasites in larger individuals, but this was found in only two locations. Contrary to expectations, we found in two localities that the intensity of parasites was lower in larger animals. The intensity was also higher in males. We did not confirm any differences in prevalence or intensity between syntopic and alotopic populations. We also did notobserve any differences related to altitude. Our results confirm an asymmetricparasitic prevalence and intensitybetween the two competing lizard species that may have a potential influence on theircompetitive relationshipand coexistence

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In parasite–host interactions host species may differ in their ability to fight parasitic infections, while other ecological interactions, including competition, may differentially alter their physiological state, making them even more susceptible to parasites. In this study, we analyse the haemogregarine blood parasites infecting two competing lizard species, Iberolacerta horvathi and Podarcis muralis, and explore host–parasite relationships under different host competition scenarios. Both species were infected with haemogregarine parasites belonging to the genus Karyolysus. Using the 18S rRNA gene, six new Karyolysus haplotypes were identified clustering with other Central and Eastern European samples, and widely shared between both lizard hosts. Haemogregarine infections were detected at all sampled sites with over 50% of individuals parasitized. Overall, I. horvathi was more frequently and also more intensely parasitized than P. muralis, with higher infection rates observed in syntopy. Males of both species tended to be more frequently infected and showed a higher infection intensity than conspecific females. The results suggest that parasitisation by haemogregarines may be relevant in the dynamics of the competitive relationship between these lizard species. More studies, including immunological response analysis, and the identification of the vectors are needed to better understand host–parasite relationships and competition.

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In parasite–host interactions host species may differ in their ability to fight parasitic infections, while other ecological interactions, including competition, may differentially alter their physiological state, making them even more susceptible to parasites. In this study, we analyse the haemogregarine blood parasites infecting two competing lizard species, Iberolacerta horvathi and Podarcis muralis, and explore host–parasite relationships under different host competition scenarios. Both species were infected with haemogregarine parasites belonging to the genus Karyolysus. Using the 18S rRNA gene, six new Karyolysus haplotypes were identified clustering with other Central and Eastern European samples, and widely shared between both lizard hosts. Haemogregarine infections were detected at all sampled sites with over 50% of individuals parasitized. Overall, I. horvathi was more frequently and also more intensely parasitized than P. muralis, with higher infection rates observed in syntopy. Males of both species tended to be more frequently infected and showed a higher infection intensity than conspecific females. The results suggest that parasitisation by haemogregarines may be relevant in the dynamics of the competitive relationship between these lizard species. More studies, including immunological response analysis, and the identification of the vectors are needed to better understand host–parasite relationships and competition.

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Parasites are important drivers shaping animal populations by affecting growth, performance, reproductive successand survival. In sympatric related host species, parasites may shape their interspecific interaction and vice-versa - the intensity of parasite infection and parasitemia couldbe influenced by beforementioned interactions. Members of the genus Hepatozoon (Apicomplexa, Adeleorina) are the most common and widely distributed intracellular parasites oflizards. In this work we studied the parasitization patterns of tw lizard species, Iberolacerta horvathi and Podarcis muralis, ranging Slovenia either in sympatry or in allopatry. We used the 18S rRNA gene to confirm parasite identity an identify potential new haplotype an estimated prevalenc and intensity using microscopy. Then, we tested for the effect of host species, geographic region, body size sex and sympatry/allopatry on these parameters. Preliminary results show significant differences in parasite prevalence and intensity among localities, as well as differences in parasite intensity both among localities and host species. This one of the first studies examining the blood parasites in Slovene reptiles combining microscop and molecular methods, and as such expands our knowledge on parasite diversity, their possible impact in this region and the interaction patterns betwee both lizard species. In addition, the study recovers the variation between sympatry and allopatry as a way of understanding coexistence of two lizard species occupying a similar ecological niche.

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This study deals with several features of the reproductive biology of Lacerta monticola cyreni. It was carried out in a lizard population in the Sierra de Guadarrama (Central Spain). Owing to the hard weather conditions the lizards have a very short active season (nearly six months). The annual gonadal cycle of both sexes shows an unimodal distribution which demonstrates the existence of only one mating and only one clutch per year. Mating occurs in May-June. Females lay a clutch of 4 to 9 eggs, usually during July. Finally, hatching takes place in September, one month before the beginning of hibernation in October.

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Das Ziel dieser Studie ist, neue Erkenntnisse über die bevorzugten ökologische Nischen und die aktuelle Verbreitung der Kroatischen Gebirgseidechse Iberolacerta horvathi (MÉHELY, 1904) innerhalb Österreichs zu gewinnen. Eine aussagekräftige Vorhersage werden wir mittels Modellierung der potentiellen Verbreitung (SDM – Species‘ distribution modeling) für Österreich treffen und darstellen können. Für die korrelative Modellierung implementieren und analysieren wir artspezifische Habitat-Präferenzen unter Berücksichtigung geologischer, topografischer, bioklimatischer und ökologischer Variablen, basierend auf Daten bisher bekannter Fundorte. Eine anschließende Freilandstudie an die vom Modell berechneten Standorte mit hoher Wahrscheinlichkeit weiterer Vorkommen, soll schlussendlich zur Entdeckung noch unbekannter besetzter Nischen von I. horvathi in Österreich führen. Endresultat der Studie wird eine umfassende Art-Charakterisierung und eine Verbreitungskarte der potentiellen, als auch realisierten Habitate, entlang der nördlichen Verbreitungsgrenze der Kroatischen Gebirgseidechse in Österreich sein. Außerdem testen wir die Aussagekraft der Modellierung, die zukünftig auch als hilfreiche, flexible Methode in Bezug auf Artenschutz Anwendung finden könnte.

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Historical factors, current population size, population connectivity and selective processes at linked loci contribute to shaping contemporary patterns of neutral genetic diversity. It is now widely acknowledged that nuclear and mitochondrial markers react differently to current demography as well as to past history, so the use of both types of markers is often advocated to gain insight on both historical and contemporary processes. We used 12 microsatellite loci genotyped in 13 populations of a mountain lizard (Iberolacerta bonnali) to test whether the historical scenario favoured by a previous mitochondrial study was also supported by nuclear markers and thereby evaluated the consequences of postglacial range movements on nuclear diversity. Congruent signals of recent history were revealed by nuclear and mitochondrial markers using an Approximate Bayesian computation approach, but contemporary patterns of mtDNA and nuclear DNA diversity were radically different. Although dispersal in this species is probably highly restricted at all spatial scales, colonization abilities have been historically good, suggesting capability for reestablishment of locally extinct populations except in fully disconnected habitats.

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A male of Aurelio’s Rock Lizard was observed devouring its own distal part of the tail after suffering an incomplete amputation (autourophagy). This is the first observed case of this phenomenon and is compared with other cannibalism observations in Sauria.

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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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Ectothermic animals living at high elevation often face interacting challenges, including temperature extremes, intense radiation, and hypoxia. While high-elevation specialists have developed strategies to withstand these constraints, the factors preventing downslope migration are not always well understood. As mean temperatures continue to rise and climate patterns become more extreme, such translocation may be a viable conservation strategy for some populations or species, yet the effects of novel conditions, such as relative hyperoxia, have not been well characterised. Our study examines the effect of downslope translocation on ectothermic thermal physiology and performance in Pyrenean rock lizards (Iberolacerta bonnali) from high elevation (2254 m above sea level). Specifically, we tested whether models of organismal performance developed from low-elevation species facing oxygen restriction (e.g., hierarchical mechanisms of thermal limitation hypothesis) can be applied to the opposite scenario, when high-elevation organisms face hyperoxia. Lizards were split into two treatment groups: one group was maintained at a high elevation (2877 m ASL) and the other group was transplanted to low elevation (432 m ASL). In support of hyperoxia representing a constraint, we found that lizards transplanted to the novel oxygen environment of low elevation exhibited decreased thermal preferences and that the thermal performance curve for sprint speed shifted, resulting in lower performance at high body temperatures. While the effects of hypoxia on thermal physiology are well-explored, few studies have examined the effects of hyperoxia in an ecological context. Our study suggests that high-elevation specialists may be hindered in such novel oxygen environments and thus constrained in their capacity for downslope migration.

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A first complete list of the reptile type specimens preserved in the Museo Nacional de Ciencias Naturales (CSIC) of Ma-drid (updated until 15 July 2012) is provided. The collection houses a total of 319 type specimens representing 24 taxa belonging to 6 families and 12 genera. There are 22 taxa represented by primary types (19 holotypes, 2 neotypes and 1 lectotype) and at least one paratype, and only two taxa are exclusively represented by one secondary type (paratype). The collection is specially rich in Spanish endemisms. Special attention is deserved by the type series of many subspecies of Podarcis lilfordi described by A. Salvador and V. Pérez-Mellado. All type specimens are housed in the Herpetological collection except Blanus mariae and Psammodromus occidentalis type series and Psammodromus hispanicus (neotype) which are preserved in the DNA/Tissues Collection.

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

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

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Identifying the factors that underlie signal divergences remains challenging in studies of animal communication. Regarding the chemical signalling, different compounds can be found in some species but be absent in others. We hypothesized that if the costs that are associated with the expression of some compounds are too high, their presence in the signal may be restricted. However, these compounds may be expressed and be functional when those costs are relaxed. Vitamin E (α-tocopherol), a dietary compound with metabolic relevancy, acts as an honest chemical sexual signal in many lizards but no in others such as the Carpetan Rock lizard (Iberolacerta cyreni). We investigated whether dietary supplementation favours the expression of this vitamin in scents of I. cyreni. We show that dietary constraints can preclude the expression of vitamin E in chemical secretions of wild males because was expressed when it was experimentally provided in the diet. Vitamin E supplementation also heightened the immune response of males and increased the interest of their scent for females, highlighting the vitamin E as a chemical sexual signal in this species. We suggest that diet could decisively act as a driver of intra- and interspecific divergences in the chemical signalling of lizards.

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Knowledge about chemical communication in some vertebrates is still relatively limited. Squamates are a glaring example of this, even when recent evidences indicate that scents are involved in social and sexual interactions. In lizards, where our understanding of chemical communication has considerably progressed in the last few years, many questions about chemical interactions remain unanswered. A potential reason for this is the inherent complexity and technical limitations that some methodologies embody when analyzing the compounds used to convey information. We provide here a straightforward procedure to analyze lizard chemical secretions based on gas chromatography coupled to mass spectrometry that uses an internal standard for the semiquantification of compounds. We compare the results of this method with those obtained by the traditional procedure of calculating relative proportions of compounds. For such purpose, we designed two experiments to investigate if these procedures allowed revealing changes in chemical secretions 1) when lizards received previously a vitamin dietary supplementation or 2) when the chemical secretions were exposed to high temperatures. Our results show that the procedure based on relative proportions is useful to describe the overall chemical profile, or changes in it, at population or species levels. On the other hand, the use of the procedure based on semiquantitative determination can be applied when the target of study is the variation in one or more particular compounds of the sample, as it has proved more accurate detecting quantitative variations in the secretions. This method would reveal new aspects produced by, for example, the effects of different physiological and climatic factors that the traditional method does not show.

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Lautäußerungen spielen eine Schlüsselrolle in einer Vielzahl von Interaktionen zwischen artgleichen und verschiedenen Individuen. Doch im Vergleich mit anderen Sinnesleistungen wird der Akustik bei Reptilien nur wenig Aufmerksamkeit geschenkt. Innerhalb der Lacertidae wird Fehlen und Vorhandensein von Lautäußerungen als euin Unterscheidungsmerkmal zwischen den Unterfamilien Gallotinae und Lacertinae angesehen, wobei letztere, höchst diverse Unterfamilie als mehrheitlich nichg stimmbegabt betrachtet wird. Mit Hilfe von Bild- und Tonaufzeichnungen berichten die Autoren erstmals über die Lautäußerungen bei Iberolacerta cyreni (MÜLLER & HELLMICH, 1937), einer europäischen Felsiedechsenart. Tonaufzeichnungen eines Exemplars im Freiland geben kurze nasale quietschende Laute wieder, die hinichtlich des Mechanismus der Schallproduktion und der akustischen Merkmale Ähnlichkeit mit den Lauten nahe verwandter Lacertidenarten zeigen. Die funktionale Bedeutung derartiger Lautgäuerungen ist unklar, doch weisen die akustischen Merkmale und der verhaltenskontect auf die Möglichkeit eines bei Gefahr abgegebenen Schreckrufs hin. Die vorliegenden Beobachtungen lassen permute, das sich die Fähigkeit zur Lautproduktion bei verschiedenen systematischen Gruppen der Halsbandeidechsen entwickelt hat und, auch wenn selten eingesetzt, weiter berbreitet ist als erwartet.

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Satellite DNAs represent a large portion of all high eukaryotic genomes. They consist of numerous very similar repeated sequences, tandemly arranged in large clusters up to 100 million base pairs in length, usually located in the heterochromatic parts of chromosomes. The biological significance of satDNAs is still under discussion, but most of their proposed functions are related to heterochromatin and/or centromere formation and function. Because information about the structure of reptilian satDNA is far from exhaustive, we present a molecular and cytogenetic characterization of two satDNA families in four lacertid species. Two families of tandemly repeated DNAs, namely TaqI and HindIII satDNAs, have been cloned and sequenced from four species belonging to the genus Iberolacerta. These satDNAs are characterized by a monomer length of 171–188 and 170–172 bp, and by an AT content of 60.5% and 58.1%, respectively. FISH experiments with TaqI satDNA probe produced bright signals in pericentromeric regions of a subset of chromosomes whereas all the centromeres were marked by HindIII probe. The results obtained in this study suggest that chromosome location and abundance of satDNAs influence the evolution of these elements, with centromeric families evolving tenfold faster than interstitial/pericentromeric ones. Such different rates render different satellites useful for phylogenetic investigation at different taxonomic ranks.

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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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The attempt is made to define the distributional patterns of the reptiles in Carinthia (Austria) by selected parameters of climat and vegetation.

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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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While global variation in taxonomic diversity is strongly linked to latitude, the extent to which morphological disparity follows geographical gradients is less well known. We estimated patterns of lineage diversification, morphological disparity and rates of phenotypic evolution in the Old World lizard family Lacertidae, which displays a nearly inverse latitudinal diversity gradient with decreasing species richness towards the tropics. We found that lacertids exhibit relatively constant rates of lineage accumulation over time, although the majority of morphological variation appears to have originated during recent divergence events, resulting in increased partitioning of disparity within subclades. Among subclades, tropical arboreal taxa exhibited the fastest rates of shape change while temperate European taxa were the slowest, resulting in an inverse relationship between latitudinal diversity and rates of phenotypic evolution. This pattern demonstrates a compelling counterexample to the ecological opportunity theory of diversification, suggesting an uncoupling of the processes generating species diversity and morphological differentiation across spatial scales.

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Mechanisms affecting consistent interindividual behavioral variation (i.e., animal personality) are of wide scientific interest. In poikilotherms, ambient temperature is one of the most important environmental factors with a direct link to a variety of fitnessrelated traits. Recent empirical evidence suggests that individual differences in boldness are linked to behavioral thermoregulation strategy in heliothermic species, as individuals are regularly exposed to predators during basking. Here, we tested for links between behavioral thermoregulation strategy, boldness, and individual state in adult males of the high-mountain Carpetan rock lizard (Iberolacerta cyreni). Principal component analysis revealed the following latent links in our data: (i) a positive relationship of activity with relative limb length and color brightness (PC1, 23% variation explained), (ii) a negative relationship of thermoregulatory precision with parasite load and risk-taking (PC2, 20.98% variation explained), and (iii) a negative relationship between preferred body temperature and relative limb length (PC3, 19.23% variation explained). We conclude that differences in boldness and behavioral thermoregulatory strategy could be explained by both stable and labile state variables. The moderate link between behavioral thermoregulatory strategy and risk-taking personality in our system is plausibly the result of differences in reproductive state of individuals or variation in ecological conditions during the breeding season.

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Vertebral malformations have been reported previously from a wide range of reptile taxa. Kyphosis refers to an abnormally excessive convex curvature of the spine in the thoracic and sacral regions, while if the deviation affects the lateral plane of the spinal column, the condition is known as scoliosis. In some cases, these two types of malformation can be combined in the same animal, known as kyphoscoliosis (Garín-Barrio et al., 2011). It appears that kyphosis mainly affects species adapted to aquatic environments (e.g. turtles and crocodiles; Tucker, 1997; Boede & Sogbe, 2000; Tucker et al., 2007), while scoliosis is more common in terrestrial species (snakes and lizards; Grogan, 1976; Simbotwe, 1983; Frutos et al., 2006). Reports of kyphosis (and kyphoscoliosis) from wild lizard populations have increased lately (e.g. Liolaemus koslowskyi (Iguandiae) Avila et al. (2013); Sceloporus torquatus (Phrynosomatidae) Pérez-Delgadillo et al. (2015); Norops sericeus (Dactyloidae) Domínguez-De la Riva and Carbajal-Márquez (2016); Sceloporus vandenburgianus (Phrynosomatidae) Valdez-Villavicencio et al., (2016); Stenocercus guentheri (Tropiduridae) Ramírez-Jaramillo, (2018); Marisora brachypoda (Scinidae) Arrivillaga & Brown,(2019).

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Behavioural consistency is expected to affect fitness in a negative way intuitively by constraining the individual behavioural repertoire. In contrast to this, consistency both within (animal personality) and across behaviours (behavioural syndrome) is observed in a wide variety of taxa. Hence, one of the main goals in behavioural ecology is to understand the evolutionary and developmental factors underlying consistent between-individual differences in behaviour. One possibility is that individual state and behaviour are linked, and thus state-behaviour feedback loops can explain the emergence of behavioural consistency. Stable state variables (i.e. life-history traits) are known to create long- lasting behavioural strategies, however, recently many research focused on the role of labile state variables in the emergence of stable behavioural differences. For instance, body condition is expected to have a strong effect on individual behavioural strategies in animals. It is known that the vitamin D component in the femoral secretion of male Carpetan rock-lizards (Iberolacerta cyreni) act as an honest sexual signal, since only males with better body condition can afford secreting vitamin D at a high rate. Hence, vitamin D and available energy are both expected to affect their behavioural consistency. In our present work, we studied the effects of food and vitamin D manipulation on the activity and risk-taking of 60 male I. cyreni during the mating season of 2014. We applied a full factorial experimental design with high vs. low food treatments and vitamin D supplementation vs. placebo treatments. We discuss the treatment effects on lizard behaviour based on eight (activity) and seven (risk-taking) repeated assays. We did not find any effect of the treatments on activity, however, our results suggests that vitamin D combined with insufficient energy intake results higher risk-taking in males compared to other treatment groups.

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Identifying evolutionary and developmental mechanisms underlying consistent between-individual differences in behaviour is the main goal in ‘animal personality studies’. Here, we explored whether activity and risk-taking varied consistently between individuals and correlated to various – potentially fitness linked – male traits in Carpetan rock lizards (Iberolacerta cyreni). Lizards showed significant consistency within both behaviours, implying the presence of activity and risk-taking personalities. However, there were no correlation between activity and risk-taking, neither on the between- nor on the within-individual levels, implying the absence of a behavioural syndrome. We found a strong link between the intensity of blood parasite (Haemogregarinidae) infection and risk-taking: lizards with higher infection intensity took more risk. While we cannot distinguish cause from causative in the parasite intensity – risk-taking correlation – our results are in line with the asset protection hypothesis predicting that individuals with lower future reproductive value should focus on the current reproductive event and take higher risk.

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It has been proposed recently that labile state variables (e.g. energy reserves) can have a key role in the development and maintenance of consistent between-individual behavioural variation (i.e. animal personality) within population. In male Carpetan rock lizards (Iberolacerta cyreni), the provitamin D3 component of femoral gland secretion acts as an honest signal in sexual communication. Further, vitamin D3 has many important metabolic functions in reptiles. Therefore, by employing a factorial experiment with food (high vs. low) and vitamin D3 (supplemented vs. control) treatments in wild-caught reproductive male I. cyreni, we tested whether changing labile components of individual state affected (i) behavioural consistency (the degree of between-individual difference) and (ii) behavioural type (mean behaviour). Animal personality in activity was present in all treatments; however, personality was present only in the high food × vitamin D3 supplementation treatment in shelter use and it was present in all but the low food × placebo treatment in risk taking. Lizards (i) decreased activity in the high food treatment, (ii) increased shelter use in the vitamin D3 supplementation treatment and (iii) increased risk taking in the low food × vitamin D3 supplementation treatment. We conclude that short-term changes in individual state affect both behavioural consistency and behavioural type of reproductive male I. cyreni. Unfavourable conditions resulted in decreased behavioural consistency, while high-state individuals became less active in general. Individuals with high specific (vitamin D3) but low general (energy reserves) state took higher risk. We discuss several evolutionary explanations for the reported patterns. Significance statement The evolutionary and developmental mechanisms resulting in consistent between-individual behavioural differences across time and situations (i.e. animal personality) are of high scientific interest. It has been recently proposed that links between individual state (e.g. how well-fed the individual is) and behaviour can maintain such between-individual differences even on an evolutionarily timescale. However, whether short-term state changes are able to affect animal personality in adults is an open question. In a manipulative experiment, we found that the amount of food and vitamin D3 (known to increase physiological quality and attractiveness of male Carpetan rock lizards, I. cyreni) affected the expression of animal personality and the actual behavioural types of reproductive male Carpetan rock lizards. Therefore, we provide evidence that short-term environmental variation does induce or suppress animal personality, and it also affects individual behaviour.

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Aggression is one of the most frequently studied behavioural traits across a wide range of taxa; however, most studies evaluate aggressive behaviour in a social context, in which aggressive interactions between conspecifics are motivated by resource control (offensive or social aggression). However, in an antipredator context, the primary role of aggression is defence (defensive or antipredator aggression). Although the neuroendocrinology of antipredator aggression is often studied in domesticated and laboratory animals, how environment and individual state affect this behavioural trait in the wild is largely unknown. Here, by conducting a manipulative experiment, we tested whether (i) consistent between-individual differences (i.e. animal personality) are present in antipredator aggression in adult male Carpetan rock lizards (Iberolacerta cyreni) and (ii) short-term environmental changes (presence vs. absence of predator cues) and differences in individual state (body length, head size, hind limb length) affect individual mean behaviour (i.e. behavioural type). We found moderate-high repeatability in antipredator aggression (willingness to bite a human), indicating the presence of animal personality in this behavioural trait. Lizards were on average more defensive in the presence of predator cues; furthermore, short-legged males showed higher antipredator aggression than long-legged males in the presence of predator cues, probably as an attempt to balance their decreased escape speed. Larger (~ older) males were more defensive than smaller ones, probably due to their increased fighting ability. We conclude that antipredator aggression is an important part of an individual’s behavioural repertoire and its expression is driven by both environmental situation and individual state.

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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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The life and work of the Alsatian Louis Amédée Lantz (1886-1953), a pioneer of European herpetology. A trained and professional research-chemist, the Alsatian Louis Amédée Lantz was an active and particularly dynamic herpetologist. His residence of nearly twelve years in Russia enabled him to make several field trips to regions such as the Caucasus and Iran. He always carried with him a home made telescopic net that he used to catch lizards. That net became symbolic of Lantz for most people around him. He realized significant herpetological collections mainly deposited in the Museum of the Zoology Institute of the Russian Academy of Sciences in St. Petersburg. Soon after the Russian Revolution in 1917, he had to leave the country and stayed for a few years in France before joining England. During this period, Lantz visited the French Pyrenees and the Mediterranean border (Hyeres Islands, Bandol), enabling him to discover for the first time oviparous populations of common lizard and also to describe a new species of lizard from the French Pyrenees. Lantz is also at the origin of the discovery of unisexual reproduction by parthenogenesis in reptiles. His collaboration with the Swedish amateur herpetologist Otto Cyrén, whom he met through the German herpetologist Willy Wolterstorff, was particularly successful throughout his career. Lantz died at the age of 66 years, in Switzerland, as he was about to join the team of herpetologists at the Natural History Museum in Paris.

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

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The International Commission on Zoological Nomenclature has conserved the names Iberolacerta and Darevskia for two genera of lacertid lizards by confirming their availability from Arribas (1999). The microfiche publication (Arribas, 1997) is confirmed as unavailable and placed on the Official Index of Rejected and Invalid Works in Zoological Nomenclature. The name Caucasilacerta Harris, Arnold & Thomas, 1998 is confirmed as a nomen nudum and placed on the Official Index of Rejected and Invalid Generic Names in Zoology.

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An updated checklist is presented for the amphibian and reptile fauna of Croatia. According to historical and recent literature sources, 20 amphibian and 41 reptile species have been recorded so far. Two reptile species, Blanus strauchi and Typhlops vermicularis, are considered doubtful, although there are individual museum vouchers for the country. Presence of a total of 59 species can be considered valid, with two additional considered doubtful. This paper summarizes data from 700 scientific and popular publications, which is the first attempt to publish a complete Croatian herpetological bibliography. Full bibliography is also given for all occurring species and constitutes a basis for future publications.

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

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the Lacertidae is one of the most diverse and widespread lizard families throughout Eurasia and Africa. Several studies so far have attempted to unravel the phylogeny of Lacertidae using morphological and mole- cular data. However, the intra-family relationships remain unclear. in an effort to explore the phylogenetic relationships within the family Lacertidae, a concatenated dataset of 5727 bp from six genes (two nuclear and four mitochondrial) and 40 genera was assembled based on GenBank database. Phylogenetic inference analy- ses were conducted using Maximum Parsimony (MP), Bayesian inference (Bi) and Maximum Likelihood (ML), revealing that even a combined dataset of both mitochondrial and nuclear genes is not able to resolve the phylogenetic relationships of the Lacertidae family under the tribe level.

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The dangers of helminth infestations for lizards, particularly infestations with nematodes und cestodes, are reported. Infection routes, diagnoses, und treatments are described.

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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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After a short synthesis of the present knowledge on the distribution of the Horvath’s rock lizard (Iberolacerta horvathi) in Italy, the Author refers about the discovery of a second population of the species on the Dolomites (Italia north-eastern Italy, Veneto Region). It dwells on the climbing wall of mount Tudaio (960 m a. s. l., Vigo di Cadore, Belluno) and has been detected thanks to the recovery of one Horvath’s rock lizard predated by a smooth snake female (Coronella austriaca) falling down from about 15 meters high. From its throat has been extracted a male of Iberolacerta horvathi determined by means of already known diagnostic characters: wide contact between rostral and frontonasal scales, flat and polygonal dorsal scales, caudal rings of regularly alternated thickness (one about a half of the other), bright white unspotted throat, bright yellow-lemon bellies. Status and consistency of the new population of Horvath’s rock lizard is not well known, but it is possible to note that at the bottom of the climbing wall the common wall lizard (Podarcis muralis) seems to be quite frequent.

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After a short synthesis of the present knowledge on the distribution of the Horvath’s rock lizard (Iberolacerta horvathi) in Italy, the Author refers about recent news and field verifications on the distribution of the lizard in Veneto and Friuli Venetia Giulia Regions (north-eastern Italy).

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After a provisional review of the knowledge on Lacerto horvathi in Italy, the authors present new data on its distribution, spacing strategy and territoriality. The species is quite common in various mountainous habitats of north-eastern Italy, both in Friuli- Venezia Giulia (Udine Province, more than 55 localities; Pordenone Province, 1 locality) and Veneto (Belluno Province, 1 locality). In these areas, it lives from 250 (Pradolino Gorge, Pulfero, Udine) to 2000 metres a.s.l.. (M.te Ponza, Tarvisio, Udine). The preliminary comparison between the homing patterns of syntopic L. horvathi and Podarcis muralis suggests that this last-mentioned species is characterised by wider home ranges — 3-9 times bigger than those of L. horvathi — showing a very high inter-specific aggressiveness. However, in deeper gorges, in cool valleys with Dinaric orientation, as well as on top of mountains, L. horvathi tends to vicariate P. muralis for various differences in both spacing, thermal, and reproductive strategies. In north-eastern Italy, such vicariance on average gradually occurs between 700 and 900 meters above sea level.

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The Authors outline the present situation and distribution maps of 118 animal species of European Community interest protected by the EU 92/43/EEC Directive in Friuli Venezia Giulia Region (North-eastern Italy). They belong to various phyla and classes (Mollusca [6 species], Annelida [1 species], Arthropoda [27 species], Vertebrata [84 species]), at various degrees of protection included in the lists of the so-called Habitat Directive (All. II, IV, V). In the present situation the main conservation problems concern various hygrophilous lowland-dwelling species, highly menaced from agriculture, pollution and habitat fragmentation. A lot of meadow-dwelling species, on the contrary, are menaced by forest expansion in various karstic and mountain environment. The great expansion of the mountain woods on the alpine mountain chain, on the other hand, has surely favoured various forest species, like a lot of big and medium Carnivores.

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Over the four year period (2007–2010), the authors conducted a field study aimed at drawing up an inventory of batrachofauna and herpetofauna of the Zrmanja river catchment area. Eight species of amphibians and twenty one species of reptiles were detected; through the available literature overview two more reptilian and one amphibian species were added to the list. Therefore, we proved the presence of thirty two species in total. High diversity of amphibians and reptiles indicates great significance of this area and imposes the need to ensure survival of these species through long-term populations monitoring and conservation measures.

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The Batuecan lizard Iberolacerta martinezricai is a critically endangered species due to ist significantly reduced distribution, which is restricted to the scree slopes (SS) of a few mountain peaks within the Batuecas-Sierra de Francia Natural Park (western Spain). Given its high specialisation in this type of discontinuous habitat, the long-term conservation of the species requires maintaining the connectivity between populations. This study analyses the contribution of connectivity, as well as other patch-related factors, in the distribution and density patterns of the species. With this aim, 67 SS were sampled by line transects from May to October 2018. Each SS was characterised using variables indicative of the microhabitat conditions for the lizard. Inter-SS connectivity was quantified using graph theory for seven distances. Generalised linear models (GLMs) were performed for both presence and density. Model results showed that while connectivity was a relevant factor in the presence of lizards, density only involved patch-related variables. Discrepancies probably occurred because the factors influencing presence operate on a wider scale than those of abundance. In view of the results, the best-connected SS, but also those where the lizard is most abundant and from which more dispersed individuals are likely to depart, seem to be the essential patches in any conservation strategy. The results may also be relevant to other species with habitat-specific requirements.

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In this paper, we present data on the female reproductive traits of the Horvath’s rock lizard from Slovenia. The clutch, egg and hatchling characteristics were investigated based on clutches laid in laboratory conditions by pregnant females collected from a natural population. A female lays one clutch of eggs annually with an average number of three (range 1-5) eggs. We found a significant positive correlation between female size and egg width and volume. The mean egg length and volume in a clutch decreased significantly with clutch size independently of female size. The incubation period averaged 44 days. Significant positive correlations were found between the hatchling total length and mass, and egg mass. There was a significant negative relationship between the egg mass and incubation duration. The life-history strategy of the Horvath’s rock lizard appeared to be shaped by several factors, such as the constraints of a high-altitude environment, flattened body morphology and possibly phylogenetic background.

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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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In spite of the importance of chemoreception and chemical signals in the social organization of lizards, there are only a few studies examining the chemical composition of secretions of lizards used for scent marking. The secretion of the femoral glands of male Iberian rock lizards (Lacerta monticola cyreni) contains 44 lipophilic compounds, including several steroids (mainly cholesterol), and n-C6 to n-C22 carboxylic acids, and minor components such as esters of carboxylic acids, alcohols, squalene, and one lactone. These compounds were identified on the basis of mass spectra, obtained by GC-MS. Most lipids were detected in all individuals, although relative proportions of each chemical show a high interindividual variability. This variability might be related to the characteristics or physical and health condition of males and might be the basis of female choice based on chemical cues observed in this lizard species.

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Multiple sexual signals may convey information on males’ quality. However, most research focused on visual signals, ignoring chemical signals. In vertebrates, chemical signals are probably often a multicomponent mixture of several active compounds, but they are not well known. We examined the potential additive and interactive effects of 2 compounds (oleic acid and ergosterol) naturally found in femoral gland secretions of male rock lizards on chemosensory exploration behavior of females. Tongue-flick (TF) rates of females to male secretions may result from the combination of responses to multiple compounds. There may be an additive or synergetic effect because male secretions with the highest proportions of both compounds received the highest TF rates, suggesting that their scents were more intriguing. However, there might be an interactive effect too; female TF rates were higher to males’ scent with high proportions of ergosterol alone, even if proportions of oleic acid were low, than to high proportions of oleic acid but with low proportions of ergosterol. Further bioassays testing TF behavior of females to standard compounds, presented alone or combined in different concentrations confirmed these findings. Variations in female TF behavior might be explained because different compounds signal different male traits of different importance for females. Our study suggested that femoral secretion of male rock lizards may act as a multicomponent chemical signal.

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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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External morphology of Horvath`s rock lizard (Lacerta liorvathi MEHELY 1904) was examined. Variability was established in the biornetric, meristic and qualitative characters, as well as differences between the adult and juvenile an males and females and between the samples of animals from the Slovenian and north Dinarie part of the area, New localities were stated and the known area of this species expanded.

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Lacerta horvathi is a relatively poorly known species assigned to a taxonomi- cally problematic group of lacertid lizards - Archaeolacerta. It is present with scattered mountain populations in the Julian Alps of Slovenia, in the Ucka Moun- tain and the northern Dinarid Mountains of Croatia, in northeastern Italy, in southwestern Austria, and in southern Germany (LAPINJ & DOLCE 1983, GRIL- LITSCH & TEDEMANN 1986, DE LUCA 1989, CAPULA & LUISELLI 1990). Much of data on distribution and taxonomy of this species were presented during the past few years (e.g. the papers cited above, DE LUCA & DULIC 1988, CAPULA et al. 1989). However, the comprehensive data regarding biology and ecology of L. horvathi are still missing. In this paper some new data on daily and seasonal activity patterns, thermoregulation, reproduction and food composition, are presented.

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Karyotype analysis of 2 male horvath`s rock lizard, Lacerta (Archaeolacerta) horvathi, found 2n = 36 in both somatic and spermatogonial metaphases. All the chromosomes are acrocentric. In diakinesis, 18 bivalents are visible. Microchromosomes, characteristic of most lacertid karyotypes, are absent in this species. In a single metaphase, satellites were observed on the 8th pair of chromosomes. Male sex chromosomes have not been determined. Overall, the karyotype of Lacerta horvathi is similar to that of Lacerta vivipara.

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Albumins from several species assigned to Lacerta s. 1. and Gallotia were investigated by means of the Micro-Complement fixation technique. The phylogenetic trees evaluated according to Fitch & Margoliash (1967) and Beverley & Wilson (1982) led us to infer a division, which is in accordance with protein electrophoretic results (Mayer & Tiedemann, 1982) as well as with the subgeneric division used before the studies of Böhme (1971) and Arnold (1973).

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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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Understanding the local effects of climate change on biodiversity is essential to guide environmental and management policies for natural areas. The lack of knowledge at the regional level has led to the development of a research program `Climate sentinels`. The main hypothesis of the program is that effects on biodiversity are particularly detectable in species that have low movement abilities. These `climate sentinel` species would be the first to respond to local climatic variations by adaptation or local extinction. From the territory of New Aquitaine (France), 20 biological indicators of climate change in different ecosystems (dune, dry, wet, mountain and forest) were chosen: plants communities of ecosystems, insects (butterflies, dragonflies, bumblebees), amphibians (Rana pyrenaica, Hyla molleri, H. arborea), reptiles (Iberolacerta bonnali, Timon lepidus, Zootoca vivipara, Emys orbicularis, Vipera berus) and mammals (Marmota marmota, micromammals). This paper presents the method of development of these indicators and the research hypotheses associated with climate sentinel species.

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Sexual selection contributes to the diversity of chemical signals in various animal groups. Lizards are good model species to study how sexual selection shapes signal diversity, as they are a chemically oriented taxonomic group with different levels of social interactions. Many lizard species bear epidermal glands secreting a waxy mixture of lipids and proteins, which are used in intraspecific communication. Previous among-species comparative analyses failed to find a relationship between the strength of sexual selection with the composition of the lipid blend in lizards. Here, we extend the investigation to the proteinaceous fraction. By using a phylogenetically informed approach, we correlated the average electrophoretic profiles of the protein from the femoral glands of 36 lacertid lizard species with the level of sexual dimorphism in size and shape, which are proxies for the strength of sexual selection. We found that as sexual size dimorphism advances, five distinct molecular weight regions in the protein profile increased their expression. Using tandem mass spectrometry, we successfully identified one of these five proteins: a carbonic anhydrase—an enzyme catalyzing the reversible hydration of carbon dioxide. Our findings suggest that proteins may be the target of sexual selection, as an active semiochemicals or as a dynamic support to other molecules: sexual selection may act indirectly on semiochemicals (namely lipids) by modifying the matrix (namely proteins).

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Individual recognition (IR) plays an important role in modulating social interactions of several animal species. IR may work at two fundamental levels: at class-level (CIR), if it allows group membership identification (e.g. familiar/unfamiliar), or at individual-level (true IR; TIR), if it allows uniquely recognizing conspecifics. Direct and indirect evidences suggest that many lizards are able to recognize conspecifics, notably using the secretions from femoral glands, specialized epidermal structures located in the cloacal region. Such secretions are made of a mix of lipids and proteins, the latter having been poorly studied but hypothesised to convey identity-related information. Using male Iberian Rock lizards, we set up bioassays to test the role of the protein fraction in IR, and specifically whether lizards: (1) can detect proteins from femoral glands, (2) can recognize their own proteins from those of an unfamiliar male (CIR) and (3) can distinguish between two unfamiliar protein signals (TIR). We found that femoral proteins can be actually detected, and the protein signal was enough to allow self-recognition, but not to distinguish between two unfamiliar males. These outcomes support the hypothesis that proteins from lizard femoral glands are used in intraspecific communication, at least at CIR level. The lack of TIR detection has three possible explanations: (1) finer IR abilities are actually absent in this species; (2) TIR requires a more complex and complete chemical signal or (3) a more prolonged and complete set of previous interactions among individuals is needed to lead to familiarity and TIR.

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The Horvath’s rock lizard (Iberolacerta horvathi) is a rupicolous mountain species endemic of the eastern Alps and northern Dinaric range. The species has its known western limit of the distribution in the Veneto region of Italy. It is not known whether the species is really rare in Veneto or whether the area has been insufficiently surveyed. In addition, it is not known whether the westward distribution of the species is limited by a physiographic or by a climatic barrier. During the period 2016-2018, 118 sites were surveyed in the Veneto and Trentino-Alto Adige regions. Four new occurrences of Iberolacerta horvathi were discovered in Veneto that: 1) largely fill the gap between the westernmost known site and the closest site to the east; 2) extend further west the known distribution by 9 km. In addition the species was confirmed in three already known sites. A species distribution model was developed with the software MaxEnt, using 100 occurrences from Italy, Austria and Slovenia. The best model shows that the distribution is explained by the asperity of their habitat, the sedimentary bedrock, the aspect, the average temperature of the coldest quarter, the rainfall seasonality and the average summer rainfall. The last variable appears as the most likely responsible for the rarefaction of the species at its western limit. In addition, the species distribution model suggest that the Horvath’s rock lizard might be present in some additional mountain groups where it has so far not been found yet

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Many reptile species dig underground nests where they deposit eggs with flexible and permeable shells that have physical contact with the soil and are highly permeable to soil water and gases. Iberian rock lizard eggs (Lacerta monticola cyrenni) incubated in acidic substrates suffered significant impairment to their development. Therefore, soil pollution could be affecting embryonic development. Low pH had a negative effect on egg water exchange, hatchling size and locomotor performance. In all cases, pH had no effect on incubation duration and embryo survival. At substrate pH of 4 and 10, eggs absorbed less water and final egg size and weight was lower than at neutral pH. Hatchlings from eggs incubated in acidic substrates had lower mass (up to 28 % of weight), SVL, and tail length than controls. Running speed – a good indicator of lizard fitness – was also affected by substrate pH. Embryos incubated at pH 4 ran slower than controls. The alteration of the water absorption process that low substrate pH had on eggs during incubation partially explained the observed effects on hatchling characteristics. These sublethal effects may influence the survival or success of juveniles during or after their first wintering.

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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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Some mate choice theories propose that only male signals that are honest and condition-dependent can be stable, while another hypothesis states that males evolve signals that exploit the sensory system of females. However, sensory traps might evolve into honest signals if they are differentially costly for males. We tested whether a pre-existing sensory bias for food chemicals explained chemosensory preferences of female Iberian rock lizards for male scents. We manipulated hunger levels of females and found that food-deprived females had increased chemosensory responses to chemical stimuli from both invertebrate prey and femoral secretions of males, but not to control water. Further tests suggested that cholesta-5,7-dien-3-ol (provitamin D3), a lipid found in both prey and males’ scent, may be one of the chemicals eliciting these responses. Moreover, hungry females spent more time on scent marks of males that had experimentally increased cholesta-5,7-dien-3-ol than on scent marks of males alone, whereas for control females this effect was not significant. We suggest that preexisting sensory bias for essential nutrients (i.e., provitamin D) may be the origin of similar female responses to male chemicals. However, previous studies have suggested that the allocation of these chemicals to ornaments is costly and only high quality males can afford it. Therefore, preexisting sensory bias for essential nutrients may further allow the evolution and maintenance of honest sexual display.

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Pheromones of vertebrates are often a mixture of several chemicals with different properties and messages, and their production seems condition dependent. Thus, pheromones are a good, but little studied, example of multiple sexual signals. Femoral gland secretions of male rock lizards Iberolacerta cyreni contain steroids that may act as pheromones, but there are also many other lipids, such as oleic acid, whose allocation to secretions may be costly because it has to be diverted from body fat reserves. This suggests that oleic acid could also have some function in secretions. Chemical analyses showed that proportions of oleic acid in femoral secretions of males were positively related to body condition of males, suggesting that the oleic acid secreted may reflect the amount of body fat reserves of a male. Tongue-flick bioassays showed that females were able to detect by chemosensory cues alone differences in proportions of oleic acid in secretions of males. Scents of males with more oleic acid elicited stronger chemosensory responses by females. Further tests with chemical standards confirmed that females distinguished oleic acid, and changes in its concentration, from other chemicals that are naturally found in secretions of males. Moreover, choice trials of scent-marked substrates showed that females were more attracted to areas that were experimentally manipulated to increase the proportion of oleic acid in natural scent marks of males. We suggest that oleic acid in femoral secretions might be a reliable advertisement of a male`s body condition, which females could use to select high-quality mates in conjunction with information provided by other chemicals. Alternatively, scent marks with more oleic acid might be simply more attractive to females if chemosensory responses of females to scent of males were originated by a preexisting sensory bias for food chemicals such as the oleic acid. Nevertheless, this sensory trap might have evolved into an honest signal because the elaboration of the signal seems differentially costly for males with different body conditions.

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Background: Many animals produce elaborated sexual signals to attract mates, among them are common chemical sexual signals (pheromones) with an attracting function. Lizards produce chemical secretions for scent marking that may have a role in sexual selection. In the laboratory, female rock lizards (Iberolacerta cyreni) prefer the scent of males with more ergosterol in their femoral secretions. However, it is not known whether the scent-marks of male rock lizards may actually attract females to male territories in the field. Methodology/Principal Findings: In the field, we added ergosterol to rocks inside the territories of male lizards, and found that this manipulation resulted in increased relative densities of females in these territories. Furthermore, a higher number of females were observed associated to males in manipulated plots, which probably increased mating opportunities for males in these areas. Conclusions/Significance: These and previous laboratory results suggest that female rock lizards may select to settle in home ranges based on the characteristics of scent-marks from conspecific males. Therefore, male rock lizards might attract more females and obtain more matings by increasing the proportion of ergosterol when scent-marking their territories. However, previous studies suggest that the allocation of ergosterol to secretions may be costly and only high quality males could afford it, thus, allowing the evolution of scent-marks as an honest sexual display.

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Scent-marked substrates may inform conspecifics on the characteristics of territorial males. Scent-marks of male Carpetan rock lizards (Iberolacerta cyreni) affect space use of females, which by selecting an area may increase the probability of mating with the male that has scent-marked that area. However, males do not hold exclusive territories, and scent-marks of different individual males are often together. This may provide complex information from multiple sources on the social structure. Here, we examined female preference in response to scent marks of various males and combinations in a laboratory experiment. Females preferred areas scent-marked by territorial old males against those scent-marked by young satellite-sneaker males. This reflected the known preference of females for mating with old males. In a second experiment, females preferred areas scent-marked by two males to areas of similar size marked by a single male. This may increase the probability of obtaining multiple copulations with different males, which may favour sperm competition and cryptic female choice, or may be a way to avoid infertile males. Finally, when we experimentally over-marked the scent-marks of an old male with scent-marks of a young male, females did not avoid, nor prefer, the over-marked area, suggesting that the quality of the old male may override the presence of a satellite male. We suggest that, irrespective of the causes underlying why a female selects a scent-marked area, this strategy may affect her reproductive success, which may have the same evolutionary consequences that “direct” mate choice decisions of other animals.

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Many lizards have diverse glands that produce chemosignals used in intraspecific communication and that can have reproductive consequences. For example, information in chemosignals of male lizards can be used in intrasexual competition to identify and assess the fighting potential or dominance status of rival males either indirectly through territorial scent-marks or during agonistic encounters. Moreover, females of several lizard species “prefer” to establish or spend more time on areas scent-marked by males with compounds signaling a better health or body condition or a higher genetic compatibility,which can have consequences for theirmating success and inter-sexual selection processes. We review here recent studies that suggest that the information content of chemosignals of lizards may be reliable because several physiological and endocrine processes would regulate the proportions of chemical compounds available for gland secretions. Because chemosignals are produced by the organismor come fromthe diet, they should reflect physiological changes, such as different hormonal levels (e.g. testosterone or corticosterone) or different health states (e.g. parasitic infections, immune response), and reflect the quality of the diet of an individual. More importantly, some compounds thatmay function as chemosignals also have other important functions in the organism (e.g. as antioxidants or regulating the immune system), so there could be trade-offs between allocating these compounds to attending physiological needs or to produce costly sexual “chemical ornaments”. All these factors may contribute to maintain chemosignals as condition-dependent sexual signals, which can inform conspecifics on the characteristics and state of the sender and allow making behavioral decisions with reproductive consequences. To understand the evolution of chemical secretions of lizards as sexual signals and their relevance in reproduction, future studies should examine what information the signals are carrying, the physiological processes that can maintain the reliability of the message and how diverse behavioral responses to chemosignals may influence reproductive success.

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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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Prey experience a broad range of temporal patterns of predation risk and may face different problems when assessing risk level and deciding how to allocate costly antipredator behaviour across time. We simulated in the laboratory repeated predatory attacks of different intensity (low versus high risk) to Iberian rock lizards, Iberolacerta cyreni, with similar numbers of low- and high-risk attacks across a trial, but with different temporal patterns of presentation (i.e. a series of low-risk and then a series of high risk, or vice versa, or successive attacks where risk changed randomly). We measured activity levels, time hidden in refuges after each predatory attack, and propensity to hide in refuges when the predator is close but not actually attacking (‘false alarm’ flights). Antipredator responses were of higher intensity when risk was high, but responses also depended on temporal patterns of risk. When previous attacks were of low-risk or when risk was random, prey activity did not change between risk levels. False alarm flights in the low-risk situation were more frequent when risk was random or immediately after a series of high-risk attacks. When successive attacks were of similar risk level, lizards seemed able to predict risk level in the next attack and modified their antipredator behaviour accurately. However, when risk level was random, lizards did not seem able to predict risk level and adopted a conservative strategy, overestimating risk even though costs of the antipredator responses increased.

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Many studies show that prey should not flee immediately from approaching predators, but should adjust their fleeing distance to minimize flight costs. We explored a new scenario where an ambush predator appears close to a relatively cryptic prey, that the predator has not yet detected. Then, instead of approaching further, the predator stops and starts looking for prey from a vantage point, while maintaining a constant distance with the still undetected prey. Probability of detection of prey will increase with time of searching. We predicted that prey should wait before escaping until the time (‘fleeing time’) when the risk of being detected, which depends on prey conspicuousness, equals fleeing costs. We tested this prediction in the field by simulating an ambush predator and examining escape decisions of rock lizards, Iberolacerta cyreni, considering two risk levels (i.e. the predator has or does not have information on the prey location) and relative conspicuousness of lizards (i.e. differences in coloration and body size). Lizards fled after some time, even if the distance between predator and prey remained constant and an attack had not been launched, probably because risk of being detected increases with time. However, to minimize fleeing costs, lizards modulated their fleeing times depending on probability of being detected (related to colour conspicuousness differences) and on size-dependent escape performance. Nevertheless, conspicuousness seemed unimportant after the predator had acquired information on the location of a prey, and/or the predator was considered as more dangerous because it had previously attacked the prey.

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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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We investigated experimentally the effect of tail loss on microhabitat use in a population of the Iberian rock lizard, Lacerta monticola . Significant differences in microhabitat use were not detected between tailed and tailless lizards during the reproductive season (spring), but during summer tailless lizards were more than often associated with rocks and used grassy and bare ground less than did tailed lizards. Conflicting requirements during reproductive versus non reproductive seasons may explain that tailless lizards do not modify microhabitat use during spring by means of behavioral flexibility.

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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 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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We describe microhabitat selection in the Iberian rock lizard Lacerta monticola, a species endemic to some high mountains of the Iberian peninsula. This species is potentially threatened, in particular, by changes in habitat conditions on the ski slopes and from associated buildings in resorts that proliferate precisely in areas it favours. We analyse the relationship between habitat physical structure and population densities, in order to establish a predictive habitat-population model. Lizards select large rock screes and Cytisus bushes, but scarcely use Juniperus bushes and grassy areas. Individuals have an aggregated spatial distribution. The application of this habitat model may allow predictions to be made about the response of L. monticola to artificial and natural habitat changes, and could help to manage suitable habitats for populations.

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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.