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

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

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

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

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

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

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

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The plastic capability of species to cope with the new conditions created by climate change is poorly understood. This is particularly relevant for organisms restricted to high elevations because they are adapted to cold temperatures and low oxygen availability. Therefore, evaluating trait plasticity of mountain specialists is fundamental to understand their vulnerability to environmental change. We transplanted mountain lizards, Iberolacerta cyreni, 800 m downhill to evaluate the plastic response in body condition, thermoregulation traits, haemoglobin level, and haemoparasite load. Initial measurements of body mass, total haemoglobin concentration ([Hb]), hematic parasite intensities, dorsal luminance, and thermoregulatory behaviour were resampled after two and four weeks of acclimation. We also tested whether an anti-parasitic drug reduced haemoparasite intensity. After only two weeks of acclimation to a lower elevation, lizards decreased 42% in [Hb], had 17% less parasite intensities, increased body condition by 25%, and raised by ~3% their mean preferred temperatures and their voluntary thermal maximum. The anti-parasitic treatment had no significant effect on the intensity of hematic parasites, but our results suggest that negative effects of haemoparasites on [Hb] are relaxed at lower elevation. The rapid plastic changes observed in thermal preferences, body condition, [Hb], and parasite intensity of I. cyreni demonstrate a potential plastic response of a mountain specialist. This may be adaptive under the climatic extremes typical of mountain habitats. However, there is uncertainty in whether the observed plasticity can also help overcome long term environmental changes.

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

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

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Sexual signals serve as an honest indicator of individual quality, reflecting either developmental and/or maintenance costs. A possible underlying physiological mechanism is oxidative stress, which could mediate energy trade-offs between sexual signals and other quality traits. In ectotherms, thermal performance acts as a key indicator of individual quality and influence signal intensity. We investigated how oxidative state is reflected in visual signals of lizards from different thermal habitats. According to our hypothesis, efficient thermoregulation requires different strategies in different thermal environments. In a habitat with predictable temperature changes, animals are less exposed to suboptimal temperature ranges and selection will, therefore, be stronger on the maximum oxidative damage at optimal body temperature. Contrarily, in a habitat with rather stochastic thermal shifts, individuals are often constricted by suboptimal thermal conditions, and oxidative damage can be limiting on a wide temperature range. We used Iberolacerta cyreni and Psammodromus algirus inhabiting stochastic and predictable thermal environments respectively. We examined two aspects of oxidative stress: the level of reactive oxygen metabolites at the preferred temperature (maximal ROM) and the temperature range in which animals produce at least 80% of the maximum level of reactive oxygen metabolites (effective ROM range). In I. cyreni, we found that duller coloration was related to a wider effective ROM range, while expression of coloration in P. algirus was negatively correlated with the maximal ROM. Our results suggest that different thermal constraints affect different aspects of oxidative damage which can indicate individual quality and are, therefore, represented in sexual ornaments.

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Interspecific competition can limit the distribution of species along altitudinal gradients. It has been suggested that Western European rock lizards (genus Iberolacerta) are restricted to mountains due to the expansion of wall lizards (Podarcis), but there is no experimental evidence to corroborate this hypothesis. This study examines if interference competition with Podarcis muralis is a plausible explanation for the alpine confinement of Iberian rock lizards Iberolacerta cyreni. In a first experiment, we used an enclosure with four types of microhabitats to investigate whether adult rock and/or wall lizards shifted microhabitat or refuge preferences in the presence of the other species, and to detect aggressive interactions between them. In a second experiment, we staged heterospecific encounters between naïve, laboratory-born juveniles to identify behavioural differences and agonistic interactions. In the enclosure, neither rock nor wall lizards changed their microhabitat preferences in the presence of the other species. Nevertheless, rock lizards increased the diversity of microhabitats and nocturnal refuges used in the single species trials, which had twice the number of conspecifics. Aggressive interactions involved mainly large rock lizard males. Juveniles did not show any interspecific agonistic behaviour, but rock lizards spent more time basking and less time moving. Thus, we found no evidence of competition between both species in terms of habitat shifts or agonistic interactions, although intraspecific interactions seemed to explain the behaviour of adult rock lizards. We conclude that other factors are currently determining the alpine confinement of rock lizards.

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West European Rock Lizards within the Iberolacerta group have a restricted distribution, with small, widely separated ranges in highland areas. The aim of this study was to identify possible habitat requirements (including habitat structure, type of vegetation, and refuge availability) and topographic factors (altitude and orientation) that may determine variations in the abundance of Iberolacerta cyreni on a 300-km2 mountain range and to discuss the implications of our results for the conservation of this endangered endemism. Both a stepwise regression and a best model selection approach showed that lizard abundance was positively correlated with only two predictors: altitude and cover of large rocks. Thus, the successful exploitation of alpine habitats by I. cyreni seemed to depend on the abundance of large rocks that may provide suitable basking substrates while minimizing predation risk. The positive association between altitude and lizard abundance predicts a fragmented distribution with isolated populations in the mountain peaks.

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Aim  In an effort to disentangle the ecological processes that confine ectotherms to alpine environments, we studied the thermoregulatory and microhabitat selection behaviours of the rock lizard Iberolacerta cyreni, which is endemic to some mountains of central Spain, and of the wall lizard Podarcis muralis, which is a potential competitor of rock lizards. Location  We chose three areas in the Sierra de Guadarrama (central Spain) that differed in their thermal quality [mean deviation of environmental operative temperatures from the lizards’ preferred thermal range (PTR)] and refuge availability: a pine forest (1770 m a.s.l.) in which P. muralis was the only species found, and two mixed shrub and rock sites (1770 and 1900 m a.s.l.) where both species were present. Methods  In the field we collected data on refuge availability, sun exposure, body temperature (Tb) and operative temperature (Te). Thus, we estimated the thermal habitat quality of the areas sampled and the thermoregulation accuracy and effectiveness of both species. Results  The pine forest had the lowest thermal quality and refuge availability. The lower-elevation shrub site offered the best thermal quality, but refuges were much scarcer than at the higher-elevation site. Both species thermoregulated accurately, because mean deviations of body temperature (Tb) from PTR were considerably smaller than those of Te. Podarcis muralis had higher Tb values than did I. cyreni, which had similar Tb values at both shrub sites, whereas P. muralis had lower Tb values at higher elevation. Overall, the thermoregulatory effectiveness (extent to which Tb values are closer to the PTR than are Te values) of both species was similar, but whereas I. cyreni thermoregulated more efficiently at higher elevation, the opposite was true for P. muralis. At the lower-elevation shrub site, I. cyreni remained closer to refuges than did P. muralis. Main conclusions  Our results suggest that the pine forest belt might prevent the expansion of rock lizards towards lower elevations as a result of its low thermal quality and scarcity of refuges, that the thermoregulatory effectiveness of rock lizards in alpine environments depends more on refuge availability than on thermal habitat quality, and that competition with wall lizards is unlikely to explain either the distribution or the thermoregulatory effectiveness of rock lizards.

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Local adaptation and range restrictions in alpine environments are central topics in biogeographic research with important implications for predicting impacts of global climate change on organisms. Temperature is strongly coupled to elevation and greatly affects life history traits of oviparous reptiles in mountain environments. Thus, species may encounter barriers for expanding their ranges if they are unable to adapt to the changing thermal conditions encountered along elevational gradients. We sought to determine whether thermal requirements for embryonic development provide a plausible explanation for elevational range limits of two species of lacertid lizards that have complementary elevational ranges in a Mediterranean mountain range (Psammodromus algirus is found at elevations below 1600 m and Iberolacerta cyreni is found at elevations above 1600 m). We combined experimental incubation of eggs in the laboratory with modelled estimates of nest temperature in the field. In both species, increasing temperature accelerated development and produced earlier hatching dates. The species associated with warmer environments (P. algirus) experienced an excessive hatching delay under the lowest incubation temperature. Moreover, newborns from eggs incubated at low temperatures showed poor body condition and very slow rates of postnatal growth. In contrast, eggs of the strictly alpine species I. cyreni exhibited shorter incubation periods than P. algirus that allowed hatching before the end of the active season even under low incubation temperatures. This was countered by lower reproductive success at higher temperatures, due to lower hatching rates and higher incidence of abnormal phenotypes. Elevational range limits of both species coincided well with threshold temperatures for deleterious effects on embryonic development. We suggest that incubation temperature is a major ecophysiological factor determining the elevational range limits of these oviparous lizards with predictable consequences for mountain distributions under future warmer climates.

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Studies on range limits clarify the factors involved in the extent of species occurrence and shed light on the limits to adaptation. We studied the effects of elevational variation on the thermal dependence of fitness-related traits (incubation time, hatching rate, and survivorship, size, and condition of hatchlings) to assess the role of incubation requirements in distribution range limits of the alpine endemic Iberolacerta cyreni. We captured gravid females from two core (summit) and two marginal (low-elevation edge) populations, we incubated their eggs at three temperatures (22, 26, and 30 °C), and we monitored phenotypic effects. Viability of eggs and hatchlings decreased, independently of elevation, as incubation temperature increased. Hatching success and embryo survivorship were lower for clutches from low-elevation areas than for those from mountain summits, showing that lizards face difficulties thriving at the low-elevation edge of their range. Such difficulties were partly counterbalanced by faster postnatal growth at lower elevations, leading to increased adult size and higher fecundity. High incubation temperature had detrimental effects also at low-elevation areas, and no elevational variation in the thermal dependence of hatchling traits was detected. We suggest that temperature effects on egg development and the lack of selective pressures strong enough to foster local adaptation at marginal areas, combined with extended egg retention, may contribute to shape the range limits of these alpine oviparous reptiles.

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

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Alpine lizards living in restricted areas might be particularly sensitive to climate change. We studied thermal biology of Iberolacerta cyreni in high mountains of central Spain. Our results suggest that I. cyreni is a cold-adapted thermal specialist and an effective thermoregulator. Among ectotherms, thermal specialists are more threatened by global warming than generalists. Alpine lizards have no chance to disperse to new suitable habitats. In addition, physiological plasticity is unlikely to keep pace with the expected rates of environmental warming. Thus, lizards might rely on their behavior in order to deal with ongoing climate warming. Plasticity of thermoregulatory behavior has been proposed to buffer the rise of environmental temperatures. Therefore, we studied the change in body and environmental temperatures, as well as their relationships, for I. cyreni between the 1980s and 2012. Air temperatures have increased more than 3.5°C and substrate temperatures have increased by 6°C in the habitat of I. cyreni over the last 25 years. However, body temperatures of lizards have increased less than 2°C in the same period, and the linear relationship between body and environmental temperatures remains similar. These results show that alpine lizards are buffering the potential impact of the increase in their environmental temperatures, most probably by means of their behavior. Body temperatures of I. cyreni are still cold enough to avoid any drop in fitness. Nonetheless, if warming continues, behavioral buffering might eventually become useless, as it would imply spending too much time in shelter, losing feeding, and mating opportunities. Eventually, if body temperature exceeds the thermal optimum in the near future, fitness would decrease abruptly.

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The ability to early detect a potential predator is essential for survival. The potential of Iberolacerta cyreni lizards to discriminate between chemical cues of their two predatory snakes Coronella austriaca (a non-venomous active-hunter saurophagous specialist) and Vipera latastei (a venomous sit-and-wait generalist) was evaluated herein. A third snake species, Natrix maura, which does not prey on lizards, was used as a pungent control. Thus, the behaviour of I. cyreni was studied regarding four treatments: (1) C. austriaca scent, (2) V. latastei scent, (3) N. maura scent and (4) odourless control. Lizards showed antipredator behaviour (such as slow-motion and tail waving) to C. austriaca and V. latastei chemicals. The antipredatory response was similar for both predators. This ability to react with an intensive behavioural pattern to the chemical cues of their predatory snakes may prevent lizards from being detected, and, if detected, dissuade the predator from beginning a pursuit.

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Cyrens’s rock lizard (Iberolacerta cyreni): a tale with a tail. The genus Iberolacerta consists of eight alpine lizard species from the Iberian Peninsula, the Pyrenees and Dinaric chanins. In this article the author focuses on one species, Cyren’s (or Carpentane) rock lizard which occurs in mountain ranges of the Eastern part of the Sistema central (Spain). Observatzions on this particular rock lizard were made in June during the years 2010-2013 in the Sierra de Gredos, and compared with data from literature. Details about distribution, behavior, reproduction and predation are presented. Special attention was given to the high frequency of autotomized tails that was observed in the adult rovck lizards.

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Die Gattung der Gebirgseidechsen (Iberolacerta) umfasst acht Arten, deren Verbreitungsgebiete auf der Iberischen Halbinsel, in den Pyrenäen sowie im Dinarischen Gebirge liegen. Dieser Artikel befasst sich mit der Zentraliberischen Gebirgseidechse, Iberolacerta cyreni, die in den Gebirgszügen im östlichen Teil der Spanischen Zentralkordillere beheimatet ist. Beobachtungen an dieser Gebirgseidechse fanden jeweils im Juni in den Jahren 2010-2013 in der Sierra de Gredos, Spanien, statt und wurden mit Angaben aus der Literatur verglichen. Es werden Details bezüglich Verbreitung, Verhalten, Fortpflanzung und Prädatoren gegeben. Besonderes Augenmerk wurde auf die hohe Anzahl autotomierter Schwänze bei den adulten Felseidechsen gelegt.

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Ultraviolet (UV) vision and UV colour patches have been reported in a wide range of taxa and are increasingly appreciated as an integral part of vertebrate visual perception and communication systems. Previous studies with Lacertidae, a lizard family with diverse and complex coloration, have revealed the existence of UV-reflecting patches that may function as social signals. However, confirmation of the signalling role of UV coloration requires demonstrating that the lizards are capable of vision in the UV waveband. Here we use a multidisciplinary approach to characterize the visual sensitivity of a diverse sample of lacertid species. Spectral transmission measurements of the ocular media show that wavelengths down to 300 nm are transmitted in all the species sampled. Four retinal oil droplet types can be identified in the lacertid retina. Two types are pigmented and two are colourless. Fluorescence microscopy reveals that a type of colourless droplet is UV-transmitting and may thus be associated with UV-sensitive cones. DNA sequencing shows that lacertids have a functional SWS1 opsin, very similar at 13 critical sites to that in the presumed ancestral vertebrate (which was UV-sensitive) and other UV-sensitive lizards. Finally, males of Podarcis muralis are capable of discriminating between two views of the same stimulus that differ only in the presence/absence of UV radiance. Taken together, these results provide convergent evidence of UV vision in lacertids, very likely by means of an independent photopigment. Moreover, the presence of four oil droplet types suggests that lacertids have a four-cone colour vision system.

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Many animals display complex colour patterns that comprise several adjacent, often contrasting colour patches. Combining patches of complementary colours increases the overall conspicuousness of the complex pattern, enhancing signal detection. Therefore, selection for conspicuousness may act not only on the design of single colour patches, but also on their combination. Contrasting long- and short-wavelength colour patches are located on the ventral and lateral surfaces of many lacertid lizards. As the combination of long- and short-wavelength-based colours generates local chromatic contrast, we hypothesized that selection may favour the co-occurrence of lateral and ventral contrasting patches, resulting in complex colour patterns that maximize the overall conspicuousness of the signal. To test this hypothesis we performed a comparative phylogenetic study using a categorical colour classification based on spectral data and descriptive information on lacertid coloration collected from the literature. Our results demonstrate that conspicuous ventral (long wavelength-based) and lateral (short wavelength-based) colour patches co-occur throughout the lacertid phylogeny more often than expected by chance, especially in the subfamily Lacertini. These results suggest that selection promotes the evolution of the complex pattern rather than the acquisition of a single conspicuous colour patch, possibly due to the increased conspicuousness caused by the combination of colours with contrasting spectral properties.

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Resources invested by prey to acquire information on predator behavior from inside the refuge are crucial to minimize the risk of suffering a future fatal attack. However, most studies have only analyzed situations where information on the predator behavior is unavailable for hiding preys. Also, temporal patterns of risk may affect antipredatory behavior allocation. We simulated in outdoor terraria series of 2 types of predatory attacks (low vs. high risks) with different sequences of risks (predictable series vs. attacks where risk level changed randomly) to Iberian rock lizards. We measured time spent entirely hidden in refuges until appearing near the exit of the refuge (appearance time) and time spent leaning out of the refuge while monitoring the predator (monitoring time) after each predatory interaction. Monitoring time, irrespectively of the temporal pattern of risk, was higher after a single low-risk approach of the predator than after a direct unsuccessful attack. In addition, after multiple repeated interactions with the predator, there was a significant decrease in monitoring time with the sequence order of interactions but only when lizards fled to the refuge after low-risk approaches. Lizards spent more time monitoring the predator from inside the refuge after a low-risk approach, which may be explained because, if there has not been a clear attack, uncertainty on immediate future risk would be greater, and prey may need more time before leaving the refuge to ensure that a predator has not detected the lizard and that it is not ambushing near the refuge. Our study suggests that acquisition of information during and after an attack is important in determining refuge use as an antipredation response.

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Animals often face situations that require quantity discrimination for decision-making. Differentiating between more and less amounts might be adaptative in different contexts such as in social relationships, navigation, or foraging. However, feeding close to conspecifics might change foraging behavior decisions due to hanges in predation risk perception and competition for resources. Here, we tested the quantity judgment abilities of foraging rock lizards (Iberolacerta cyreni) in a spontaneous choice test between 5 and 10 cricket prey. Lizards tested had been previously housed alone or in a group to examine how social experience affected foraging decisions. Our results showed that “grouped” lizards had greater foraging activity than “solitary” lizards. Furthermore, all lizards demonstrated some ability to discriminate between the two amounts of prey. These results reveal the capability of quantity judgments in rock lizards and also that foraging decisions may be modulated by the previous social experience.

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An essential part of foraging ecology is to understand the processes of detection, recognition and discriminationof prey, as well as the sensorial modalities involved. Often, predators do not rely on a single sensory system but on multipleinteracting senses. Specifically, lizards mainly use vision and vomerolfaction for prey pursuit. Here, we used an experimentalapproach to study how the Carpetan rock lizard,Iberolacerta cyreni, responds to different types of stimuli (chemical, visual,or both combined) from two prey species. The number of individuals approaching the prey and the number of attacks differedbetween treatments, however, we did not find differences in latency time, number of individuals attacking the prey or numberof tongue flicks. Our results suggested that visual cues combined with chemical stimuli enhanced detection of both preyspecies and that prey discrimination occurred posteriorly and independently of using any or both types of stimuli.

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A survey of the helminth communities of a population of Iberolacerta cyreni (MÜller et Hellmich, 1937), a small lizard endemic to Sistema Central (Iberian Peninsula), was conducted to determine the prevalence, abundance and species diversity of parasites of these reptiles. Four species of helminths were found, one trematode, Plagiorchis molini Lent et Freitas, 1940, one cestode, Nematotaenia tarentolae López-Neyra, 1944, and two nematodes, Skrjabinelazia sp. and Spauligodon carbonelli Roca & García-Adell, 1988. Helminth infracommunities of I. cireny showed very low values of abundance and species richness and diversity, being similar to other European lizards. The parasites found seem to have no influence on the conservation status of the host species in its natural habitat.

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There is growing concern that anthropogenic climate change may particularly affect ectotherm groups such as amphibians and reptiles. Effects of anthropogenic climate change were already observed on different levels, ranging from changes in the phenology of individuals to changes in entire populations, their demography and geographic ranges with losses and range gains. Especially the latter is exacerbated by interactions of climate change with existing risk factors such as the spread of patho- gens and invasive species. In extreme cases, species are threatened by extinction, which is most likely in endemic species occupying restricted ranges. In this review, possible future climate change scenarios as well as previous observations and general expectations of climate change induced changes in phenology, demography and bio- geography of amphibian and reptile species are presented. Individual case studies on climate change induced changes in demography of the common lizard as well as possible range shifts in the alpine newt are highlighted, with applied methods being discussed regarding different elements of uncertainty.

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Female Carpetan rock lizards (Iberolacerta cyreni) might assess the quality of males from their chemical signals. Females select areas scent marked by males with secretions containing high proportions of provitamin D to ensure mating with high-quality males. However, an alternative explanation might be that females are not choosing a mate, but that females have a pre-sensory bias for chemical cues of vitamin D in the food and are, in fact, assessing habitat quality to obtain direct benefits of increasing the intake of vitamin D. We evaluated experimentally the possible benefits of a nutritional supplement of provitamin D or vitamin D in pregnant females for their clutches and offspring. However, we did not find large differences between treatments, except in the lower body condition of juveniles of mothers supplemented with provitamin D. We also tested the chemosensory interest of females in the scent of males to study the existence of a sensory bias; this response decreased when the amount of dietary vitamin D exceeded their needs. The results suggest that there are no reproductive benefits in the intake of additional vitamin D and that the potential sensory bias to scents of males could be related to the physiological needs of the females.

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One way that maternal effects may benefit the offspring is by informing them about the characteristics of the environment. Through gestation, environmentally induced maternal effects might promote in the offspring-specific behavioral responses like dispersal or residence according to their new habitat characteristics. Females of the Carpetan rock lizard (Iberolacerta cyreni) seem to choose their home ranges using the smell of provitamin D3 in scent marks produced by males. Here, we supplemented gravid females of I. cyreni with dietary provitamin D3 or vitamin D3 to examine whether these food resources, also associated with the scent of males, affect the motivation to disperse and the locomotor performance of their offspring. Our results suggest that the supplementary availability of the resource (vitamin D3) to mothers may provoke condition-transfer maternal effects that motivate the residence or the dispersal of the offspring in their postnatal habitat. Thus, hatchlings of supplemented females had a lower dispersal trend in spite of having a greater climbing ability than hatchlings from nonsupplemented females. This suggests that the levels of provitamin D3 and vitamin D3 inside the body of the mother could act as an informative compound of the habitat quality for the offspring.

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Sexual signals can be evolutionarily stable if they are condition dependent or costly to the signaler. One of these costs may be the trade-off between maintaining the immune system and the elaboration of ornaments. Experimental immune challenges in captivity show a reduction in the expression of sexual signals, but it is not clear whether these detrimental effects are important in nature and, more importantly, whether they have reproductive consequences. We designed a field experiment to challenge the immune system of wild male Carpetan rock lizards, Iberolacerta cyreni, with a bacterial antigen (lipopolysaccharide). The immune challenge decreased relative reflectance of ultraviolet structural and melanin-dependent sexual coloration in the throat and the lateral ocelli, whereas the carotenoid-dependent dorsal green coloration was not affected. Immune activation also decreased proportions of ergosterol and cholesta-5,7-dien-3-ol in femoral secretions. These results support a trade-off between the immune system and both visual and chemical sexual ornaments. Moreover, the reproductive success of males, estimated with DNA microsatellites, depended on the expression of some color and chemical traits. However, the immune challenge did not cause overall differences in reproductive success, although it increased with body size/age in control but not in challenged males. This suggests the use of alternative reproductive strategies (e.g., forced matings) in challenged males, particularly in smaller ones. These males might consider that their survival probabilities are low and increase reproductive effort as a form of terminal investment in spite of their “low-quality” sexual signals and potential survival costs.

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Foraging strategies aim to maximize the amount of food obtained while minimizing searching costs. To reduce these costs, animals use different strategies based on the use of personal or social information to exploit food patches. At the same time, the social attraction for food resources could increase competition intensity for them. Prior experiences of animals regarding social risk and the foreknowledge of the competitors might drive the foraging strategies. In this paper, we examined experimentally whether rock lizards used behavioural strategies to reduce the risks of foraging in presence of potential competitors. We measured the foraging behaviour of a lizard resident to a territory (i.e. terrarium), in the presence of both familiar and unfamiliar conspecifics (po tential competitors). We considered whether foraging choices between two food sources of different value (i.e. quantity) are influenced by familiarity with the intruder and the evaluation of its competitive ability based on body size differences between lizards. We found differences in the number of attacks performed to the best food source, with more attacks when the intruder was unfamiliar. The results suggest evidence of both dear enemy recognition and current rival assessment modulate the foraging choices depending on the identity and the social relationship with the intruder.

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

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We followed a field population of the alpine lizard Iberolacerta cyreni over 2 consecutive breeding seasons and assigned paternity to the offpring using 8 microsatellite markers. Paternity data, combined with observations of the behavior, morphology, and spacing patterns of lizards, allowed us to document the extent of polygamy, the phenotypic correlates of the number of offspring sired, and the correlation between male reproductive success (RS) and probability of recapture the second year of our study. Multiple paternity was observed in nearly 50% of clutches, and the mating system was highly polygynandrous. In the first year of our study, male RS increased with body size, activity, tail length, and color saturation of the dorsum. In the second year, male RS increased with activity and body condition. Overall, increased male activity, a trait that is expected to decrease survivorship, was the explanatory variable that had the greatest effect on RS. However, the residents of our first study year that were recaptured in the second year had longer tails, were more active, and sired more offspring than their conspecifics that were not recaptured. Thus, contrary to expectations, no negative correlation between present reproduction and future survival was found, which suggests that male investment in reproduction is condition dependent and positively correlated with the ability to pay the underlying costs of increased activity.

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

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

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

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Here, we report the results of a large-scale PCR survey on the prevalence and diversity of adenoviruses (AdVs) in samples collected randomly from free-living reptiles. On the territories of the Guadarrama Mountains National Park in Central Spain and of the Chafarinas Islands in North Africa, cloacal swabs were taken from 318 specimens of eight native species representing five squamate reptilian families. The healthy-looking animals had been captured temporarily for physiological and ethological examinations, after which they were released. We found 22 AdV-positive samples in representatives of three species, all from Central Spain. Sequence analysis of the PCR products revealed the existence of three hitherto unknown AdVs in 11 Carpetane rock lizards (Iberolacerta cyreni), nine Iberian worm lizards (Blanus cinereus), and two Iberian green lizards (Lacerta schreiberi), respectively. Phylogeny inference showed every novel putative virus to be a member of the genus Atadenovirus. This is the very first description of the occurrence of AdVs in amphisbaenian and lacertid hosts. Unlike all squamate atadenoviruses examined previously, two of the novel putative AdVs had A+T rich DNA, a feature generally deemed to mirror previous host switch events. Our results shed new light on the diversity and evolution of atadenoviruses.

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The authors submit a supplement to the 1994 catalogues of type specimens of the herpetological collection at the Natural History Museum, Vienna (Austria). Additions comprise amphibian and reptile type specimens designated after January 1st, 1994 as well as specimens which were not recognized as types earlier.

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

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