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of the real lizards, family Lacertidae
Iberolacerta ARRIBAS, 1997
Aguado de la Paz, S. (2016) -
Las restricciones térmicas pueden limitar la fisiología y el comportamiento de los ectotermos debido a la dependencia térmica de sus funciones metabólicas. Los animales termorreguladores han desarrollado mecanismos de ajuste, tanto comportamentales como fisiológicos, que permiten mantener sus temperaturas corporales en el rango de temperaturas óptimas para la mayoría de las funciones fisiológicas. Además, ciertos rasgos morfológicos pueden favorecer el intercambio de energía con el ambiente así, un menor tamaño corporal aumenta las tasas de calentamiento, del mismo modo que los colores oscuros, por su mayor capacidad de captación de la radiación. En esta tesis, se analiza la termorregulación globalmente, manejando varias especies de saurios para dilucidar los mecanismos que optimizan el proceso de termorregulación en los ambientes más desfavorables. Utilizando como modelo una especie de alta montaña, Iberolacerta cyreni, se estudian los mecanismos de termorregulación empleados en ambientes óptimos y sub-óptimos, explorando el papel del tamaño corporal como rasgo morfológico determinante en la biología térmica. En segundo lugar, se establece el efecto de la temperatura sobre el crecimiento juvenil temprano, rasgo relevante por su influencia sobre el tamaño corporal en la madurez, comparando dos poblaciones de ambientes térmicos contrastados en Iberolacerta monticola. El estudio del rol del melanismo en la biología térmica se aborda desde dos perspectivas. Primero, utilizando como modelo la especie I. monticola, se comparan los patrones de coloración en dos poblaciones de los extremos de su distribución altitudinal, analizando simultáneamente las diferencias locales en la cantidad de melanina y comparando los valores actuales con los de especímenes preservados recolectados en las mismas poblaciones en 1983-1984, discutiendo los posibles efectos del cambio climático. Además, se determina el papel del melanismo sobre las tasas de calentamiento bajo distintas condiciones térmicas. Por último, se midió la variación en la reflectancia y el tamaño corporal, así como en respuesta térmica en la especie Cordylus cordylus, endémica de Sudáfrica. Analizando los resultados bajo la hipótesis del melanismo térmico, se evalúa la hipótesis de la co-adaptación entre la fisiología térmica, el tamaño corporal y el melanismo, valorando la influencia de otros factores de selección, como la cripsis o la disponibilidad de refugios adecuados. Nuestros resultados indican que I. cyreni es una especie termorreguladora activa bastante precisa pese a las limitaciones de las temperaturas disponibles en el campo, mostrando temperaturas preferidas inferiores a la mayoría de lacértidos. El mantenimiento de la temperatura corporal se realiza fundamentalmente mediante ajustes comportamentales, incluso en ambientes sub-óptimos con elevados costes asociados. Por otro lado, en I. monticola, existen diferencias locales en el crecimiento temprano en masa independientes del ambiente térmico, mientras que la norma de reacción del crecimiento en longitud mostró patrones opuestos entre poblaciones. Además, se encontraron evidencias de crecimiento compensatorio, con patrones opuestos en la interacción entre la población y el ambiente térmico entre las poblaciones. El grado de melanismo en I. monticola varía dependiendo de la población (altitud), el sexo y el periodo temporal (1983-1984 vs 2011-2012), con lagartijas más oscuras a mayor altitud, machos más oscuros que hembras, y mayor dimorfismo sexual en la población de montaña en la muestra actual (2011-2012). Este patrón complejo de variación sugiere la existencia de diferentes procesos y mecanismos causales que actúan sobre la evolución de la coloración. Por último, los resultados muestran que en C. cordylus, el tamaño corporal, el melanismo y las adaptaciones térmicas podrían estar co-adaptadas, aunque otros factores, como la selección sexual y la cripsis podrían intervenir en la variación del melanismo entre poblaciones.
Aguado, S. & Brana, F. (2014) -
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.
Akeret, B. (2017) -
Alagic, A. & Zagart, A. & Krofel, M. & Lazic, M. (2018) -
We analysed asymmetry in the common wall lizard (Podarcis muralis) and the Horvath`s rock lizard (Iberolacerta horvathi) from 16 populations in Slovenia and Croatia to understand their connection with potential stress factors: altitude, urbanization and the presence of another species (interspecific competition). We also compared morphological body characteristics between the two species. We used geometric morphometric methods to create landmark coordinates in the photographs of lizards` heads. Modified ANOVAs and an asymmetry index were used to find the differences in four physical characteristics: shape and size of the head, body size and the number of supraciliar scales. We found fluctuating asymmetry in the number of supraciliar scales and the shape of the head in all of our samples. Both species had more asymmetric heads in the intermediate altitudes, which might be connected with lower availability of suitable habitats. Common wall lizards from urban environment did not have more asymmetries compared to those living in natural environments. Contrary to our expectations, we found more asymmetries in allotopic than syntopic populations, indicating limited effect of interspecific interactions. Horvath`s rock lizards had more asymmetric heads, which suggests they might be exposed to greater environmental stress, although genetic effects are also possible, since directional asymmetry was also detected.
Almeida, A.P. & Rosa, H.D. & Paulo, O.S. & Crespo, E.G. (2002) -
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.
Alves de Matos, A.P. & Caeiro, M.F. & Vale, F.F. & Crespo, E. & Paperna, I. (2013) -
Icosahedral nucleo-cytoplasmic large DNA viruses (NCLDV)-like viruses, which form inclusions in the erythrocyte cytoplasm of reptiles, were previously presented as candidates for a new genus of the Iridoviridae family. The present work describes the distribution of infected lizard hosts and ultrastructural characteristics of the viral inclusions of NCLDV-like viruses from Portugal and adjacent locations in Spain. Giemsa-stained blood smears of 235 Lacerta schreiberi from Portugal and Spain, 571 Lacerta monticola from the mountain Serra da Estrela (Portugal), 794 Podarcis hispanica from several localities in Portugal and Spain, and 25 Lacerta dugesii from Madeira Island, were studied. Infection in L. schreiberi was only found in mountain populations, up to 30% in Serra da Estrela and 9–11% elsewhere. It was absent in lizards from lowlands. Prevalence of infection among L. monticola in Serra da Estrela was 10%; infected lizards were found during March to July and October but not in August and September. Infection in P. hispanica was below 3.3%. Only one infected specimen of L. dugesii was identified by light microscopy. Ultrastructural examination of infected samples revealed that the inclusions are virus assembly sites of icosahedral cytoplasmic iridovirus-like virions. Virions from different host species have different ultrastructural features and probably represent different related viruses.
Alves de Matos, A.P. & Crespo, E.G. & Paperna, I. (1998) -
Alves de Matos, A.P., Paperna, I, Rosa, H.D., Sa Sousa, P. (1994) -
Amat Orriols, F. (2003) -
Amat, F. & Pérez-Mellado, V. & Hérnandez-Estévez, J.& Garcia-Diez, T. (2008) -
We report the first study of diet composition of a Pyrenean lizard of the genus Iberolacerta, the Aurelio`s lizard, I. aurelioi, living in an alpine rocky slope at 2300 m of altitude in Andorra. Diet composition was studied during two years from 289 faecal pellets containing 966 prey items. Number of prey per pellet shows annual, seasonal and sexual differences. Diet of the species is poorly diverse, mainly dominated by insects, but we find clear evidences of opportunistic cannibalism based on the presence of toes and scales in feces. In addition, we detected minor sexual, annual and month differences in prey presence and abundance. Low amplitude of trophic niche width and prey diversity, as well as predation on clumped prey and cannibalistic behaviour suggest that Iberolacerta aurelioi has a remarkable dietary strategy in response to strong thermal and food availability constraints at high mountain habitats.
Amo, L. & López, P. & Martin, J. (2004) -
The study of the effect of parasites on their host populations is essential for understanding their role in host population dynamics and ecology. We describe the general field population biology of haemogregarines in the Iberian rock lizard, Lacerta monticola, examining the factors that determine the prevalence and intensity of infection. Prevalence and infection intensity were higher in adults than in juvenile lizards. The prevalence rate was higher in larger lizards, probably because they were also older and had been more often in contact with parasites during their lifetime. During the mating season, the intensity of parasite infection was greater in males than in females, probably due to immunosuppressive effects of testosterone. The parasite load had a negative effect on the body condition during the reproductive season. The results suggest that the interactions between parasites and hosts are not stable in this lizard population.
Amo, L. & Lopez, P. & Martin, J. (2007) -
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.
Amo, L. & López, P. & Martín, J. (2007) -
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.
Amo, L. & López, P. & Martin, J. (2007) -
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.
Andrada, J. (1980) -
Angel, F. (1946) -
Anonymous (2003) -
Antunes, P. & Crespo, E.G. & Vicente, L. (2001) -
Previously known in Portugal only in Serra da Estrela (Centre of Portugal), the Iberian rock-li zard, Lacerta monticola, is now reported inside the Montesinho Natural Park (North of Portugal). The specimens found in Museu Bocage`s collection could be tentatively assigned to subspecies call1abrica.
Antunes, P. & Vicente, L. (2004) -
The last catalogue of the Reptile collection of the Museu Bocage dates back to 1972. The collections were lost in a fire 25 years ago, so it was necessary to analyse the effort of various donors, who offered the Museum a new collection. The evaluation of the sampling effort is important to gather information (on quantity and origin) and the various samples of the species of lacertid lizards present in the Museum and enables an operational and practical visual imaged of that information. The obtained results prove that the study of the sampling effort using ordinary kriging method is robust and accurate on identifying gaps (under or no sampled areas) and a helpful tool to fulfil them. Although beginning to the representative of the Portuguese fauna, the lacertid collection of the Museu Bocage needs major improvements given that it still does not illustrate the actual distribution of lacertid lizards in Portugal. In order to stand for modern standards, when complementing the existing collection we must take into account not only the information provided by the analysis but also conservationist concerns (not killing animals) and the establishment of co-partnerships with congener institutions as well. Nowadays Museums have the duty, more than ever, to keep their collections well housed, cared for and accessible to the scientific community.
Aragón Carrera, P. (2001) -
Aragón, P. & López, P. & Martin, J. (2000) -
Aragón, P. & López, P. & Martin, J. (2001) -
We conducted a field study to examine the significance of the seasonal changes in activity levels, spatial relationships, and social interactions of the Iberian rock lizard, Lacerta monticola, during the same favorable climatic period. Activity level, movement rate, home-range size, number of same-sex neighbors, and number of agonistic interactions decreased as the mating season finished for males but not for females. The number of females` home ranges overlapped by the home range of each male was also higher in the mating season than in the nonmating season. Measurements of available ambient temperatures showed that the thermal environment was suitable for the activity of lizards in both seasons, therefore the seasonal changes cannot be explained by thermal constraints. Males gain access to several females by increasing their home-range size, thus overlapping a larger number of females` home ranges, but as a consequence, also those of a larger number of male competitors. Thus, the cost of incurring agonistic interactions also increases. These results suggest that because activity and aggressiveness in males are related to their mating success, the balance between the costs and benefits of their activity and spatial strategies differs seasonally, and this may be the cause of the observed seasonal changes.
The ability of territorial lizards to discriminate between scents of neighbors and non-neighbors might contribute to decreasing the costs of aggressive interactions. To test this hypothesis, we conducted a field study to analyze the spatial relationships between male Iberian rock-lizards, Lacerta monticola. We then used the same individuals in a laboratory experiment to test whether male lizards can use chemical cues to discriminate between familiar conspecific males (those whose home ranges overlapped) and unfamiliar conspecific males (those whose home ranges did not overlap, and whose home range centers were at least 50 m apart). Differences in tongue-flick rates in the presence of chemical cues suggested that male L. monticola discriminated between odors of familiar and unfamiliar males. The behavioral responses were also dependent on relative differences in body size between the responding male and the unfamiliar male that donated the scent: There was a significant negative correlation between tongue-flick rates emitted in cages of unfamiliar males and the body size differences between males. In contrast, when the donor of the scent was a familiar male, the tongue-flick rate was not dependent on body size differences. These results are compatible with individual discrimination through chemical cues in male L. monticola.
Aragón, P. & López, P. & Martin, J. (2004) -
We conducted a field study to analyze spatial relationships among male Iberian Rock Lizards, Lacerta monticola. We then used the same individuals in a laboratory experiment to test whether avoidance responses of a male lizard in a previously unknown area is affected by presence of scents of familiar and unfamiliar resident males. Time spent attempting to escape in presence of unfamiliar odors was significantly higher than in presence of familiar odors suggesting avoidance of unfamiliar scent marks, which may lower the costs of aggressive interactions. Our results are concordant with previous studies in which differential tongue-flick rates showed discrimination between odors of familiar and unfamiliar males of L. monticola.
Aragón, P. & López, P. & Martin, J. (2006) -
Many studies on contest competition used residency
asymmetry as a discrete variable. However, the
probability of winning an interaction may change as a
continuous function of the value of the location where the
encounter occurs. We performed a field study to examine
the importance of location within a home range and
relative body size to the outcomes of agonistic interactions
between male lizards, Lacerta monticola. The distances to
activity centers (themost used locations based on a density
function of sightings) and relative size play important
roles in agonistic interactions and had interacting effects in
natural conditions. On the other hand, previous studies
with lizards suggested that inferior competitors are able to
avoid agonistic interactions in the field. Thus, we staged
encounters in the laboratory to examine the behavioral
responses of smaller individuals. The responses of each
focal smaller male were measured in its own home cage
(resident), in the cage of a larger male (intruder) and in a
cage in which no male was previously present (control).
The predominant behavioral tactics of smaller males were
avoidance when they are the intruders and displaying
when they are the residents. Submissive displays by
smaller males may help reduce the costs of agonistic
Aragón, P. & López, P. & Martin, J. (2007) -
Theoretical and field studies suggest that inferior competitors may be able to avoid costly agonistic interactions, although the mechanisms involved are poorly understood in lizards. The “Dear enemy recognition” theory proposes that higher tolerance among familiar than among unfamiliar conspecifics may reduce costly interactions. Male lizards Lacerta monticola are able to discriminate between familiar and unfamiliar conspecific males through chemical signals alone. To better understand the implications of this discrimination ability, we performed a laboratory experiment on the role of familiarity in social tolerance estimated as the degree of spatial proximity with no sign of aggressiveness between males with a size asymmetry. In natural conditions the outcome of neighbour-stranger interactions might be the result of asymmetries in residence. Therefore, we staged pairwise encounters between familiar and unfamiliar males in neutral terraria to avoid an interacting effect of residency asymmetry. The latency to the first time in close proximity was significantly longer when small males were faced with unfamiliar males. The time that unfamiliar lizards spent in close proximity was significantly lower than that with familiar males. Our design allowed us to conclude that there was a higher social tolerance among familiar individuals with a size asymmetry that was independent of residency asymmetry.
Aragón, P. & López, P. & Martín, J. (2008) -
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.
Argüello, J.A. (1990) -
Argüello, J.A. & Salvador, A. (1988) -
Activity, temperature and habitat selection of Lacerta monticola were studies at one locality in the Cantabrian mountains. L. monticola has an unimodal pattern and is relatively inactive after midday. There is no significant different in habitat use between males, females and subadults. Lizards do not seasonally vary their patterns of habitat utilization. There is a significant correlation between body temperature and ambient temperature. Also, body temperature is significantly correlated with substrate temperature. Males, females and subadults did not differ significantly in body temperature. Body temperatures varies with time of day but not among months.
Aribas, O.J. (2008) -
Arnold, E.N. (1984) -
The ability to shed (autotomize) all or part of the tail, usually in response to predator attack, and often to subsequently regenerate it is widespread in lizards and amphisbaenians and also occurs in a few snakes and in the tuatara. Most species possess a sophisticated intravertebral autotomy mechanism which seems to be primitive in the Squamata. This appears to have been independently lost in members of many groups, but some agamids and snakes have regained the ability to shed their tails by a simpler intervertebral means and a -number of agamids have also redeveloped tail regeneration as well. Breakable tails are used to evade capture in two main ways: by enabling reptiles to break away from predators that have grasped them by the tail and by providing a distraction which deflects the attention of the attacker away from the vulnerable head and body. It is argued that loss of caudal autotomy has occurred when the costs of tail shedding outweigh its benefits. Likely costs include the expense of regrowing the tail and the loss of a variety of possible tail functions that may cause partial incapacitation, at least until the tail regenerates. Benefits of autotomy are liable to be low if predation is rare, if the animal is able to protect itself effectively in other ways, if it is too slow to evade further pursuit after the tail is shed, or if the tail is small or unpalatable and consequently not likely to distract a predator. Benefit variation may well be greater than cost variation and therefore more important in initiating the loss of autotomy mechanisms. Many taxa that do not shed the tail appear to conform to the above interpretation, but in some cases, such as the Platynota, Agamidae and Chamaeleonidae, lack of intravertebral autotomy may reflect the history of these groups rather than being a direct result of present ecological pressures. The distribution of intervertebral autotomy in the Agamidae suggests that it may have evolved only in rather special circumstances where tail fragility is advantageous even in the absence of the ability to regenerate. Restriction ot autotomy planes to the tail-base, so that the whole organ is lost, a condition found in a number of relatively slow-moving geckoes, is interpreted as a means of ensuring that enough of the tail is shed to distract a predator from further pursuit. The stimulus necessary to induce autotomy can vary rapidly in individual lizards and at least some of these changes probably maximize the effectiveness of the tail-shedding mechanism. Differences in the readiness with which all or part of the tail is shed exist between species and are likely to reflect the balance of costs and benefits in particular cases. Variations in incidence of broken tails between species and populations may be due to such differences in fragility but many other factors may play a part, including the age structure of samples, incidence of unsuccessful attacks by predators and ability to evade predators after autotomy. There is a clear tendency for climbing lizards, especially those living on rock surfaces, to have higher incidences of broken tails than ground-dwelling species, perhaps because the tail is usually less important in locomotion in the first group. Many lizards possess conspicuously coloured tails and tail movements that seem likely to help distract attention from the head and body. Conspicuous tail colouring is more frequent and often better developed in young animals, which tend to be more vulnerable than adults, and in active species from open habitats where crypsis may not always be very effective. Conspicuous tails usually have contrasting light and dark areas in nocturnal forms but are often a single bright colour in diurnal ones, probably reflecting the visual capacities of their respective predators. The predominance of blue tails in day-active species may be because this colour is striking close to but not very arresting at a distance, so it may not attract predators from far away while still drawing their attention at close quarters.
Arnold, E.N. (1987) -
Twenty-four species of lacertid lizards were examined at 31 sites in western Yogoslavia, Greece and Iberia. Comparative observations were made on over 4500 individual lizards, noting such features as times of activitiy, hunting methods, diet, micro- and macrohabitat, refuges used and body temperature. These data are used as a basis for assessing resource partition in related sympatric species, for find out whether different systematic groups have characteristic types of niche, and for comparing cumminity structures in the peninsulas of southern Europa.
Acanthodactylus erythrurus, Algyroides marchi, Algyroides moreoticus, Algyroides nigropunctatus, Lacerta agilis, Lacerta graeca, Lacerta horvathi, Lacerta lepida, Lacerta monticola, Lacerta oxycephala, Lacerta mosorensis, Lacerta schreiberi, Lacerta trilineata, Lacerta viridis, Lacerta vivipara, Podarcis bocagei, Podarcis erhardii, Podarcis hispanica, Podarcis melisellensis, Podarcis milensis, Podarcis muralis, Podarcis peloponnesiaca, Podarcis sicula, Podarcis taurica, Psammodromus algirus, Psammodromus hispanicus.
Arnold, E.N. (2002) -
Differences in surface structure (ober- hautchen) of body scales of lacertid lizards involve cell size, shape and surface profile, presence or absence of fine pitting, form of cell margins, and the occurrence of longitudinal ridges and pustular projections. Phylogenetic information indicates that the primitive pattern involved narrow strap-shaped cells, with low posteriorly overlapping edges and relatively smooth surfaces. Deviations from this condition produce a more sculptured surface and have developed many times, although subsequent overt reversals are uncommon. Like variations in scale shape, different patterns of dorsal body microornamentation appear to confer different and conflicting performance advantages. The primitive pattern may reduce friction during locomotion and also enhances dirt shedding, especially in ground-dwelling forms from moist habitats. However, this smooth microornamentation generates shine that may compromise cryptic coloration, especially when scales are large. Many derived features show correlation with such large scales and appear to suppress shine. They occur most frequently in forms from dry habitats or forms that climb in vegetation away from the ground, situations where dirt adhesion is less of a problem. Microornamentation differences involving other parts of the body and other squamate groups tend to corroborate this functional interpretation. Microornamentation features can develop on lineages in different orders and appear to act additively in reducing shine. In some cases different combinations may be optimal solutions in particular environments, but lineage effects, such as limited reversibility and different developmental proclivities, may also be important in their genesis. The fine pits often found on cell surfaces are unconnected with shine reduction, as they are smaller than the wavelengths of most visible light.
Arnold, E.N. & Arribas, O. & Carranza, S. (2007) -
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.
Arribas, O. (2000) -
Arribas, O.J. (1993) -
Lacerta monticola bonnali Lantz, 1927, that inhabits central Pyrenees from 1.800 m to ca. 3.000 m a.s.l. in alpine and subalpine environments, should be considered hereinafter as a full species: L. bonnali stat. nov. due to strong differences in pholidosis, design and coloration, as well as ecological (types of rocks among it lives) and biological features (egg incubation time) that distinguish it both from L. monticola Boulenger, 1905 and L. horvathi Mehely, 1904, their closest geographic relatives, as well as from other Archeolacerta from Europe and Near East.
Based on the study of geographic variability (pholidosis and biometry) of the Pyrenean endemite Lac
bonnali LANTZ, 1927, a new subspecies of this lizard is described from Vali d`Aràn and Pallars.
Arribas, O.J. (1994) -
We give an historical review of the discovery and knowledge of A. bonnali in the
Pyrenees and Catalonia. The up till now known locatities, habitat and specimens
morphology are included.
Arribas, O.J. (1995) -
Arribas, O.J. (1996) -
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.
Arribas, O.J. (1997) -
The geographic limits of the three high mountain pyrenean lizards (Archaeolacerta bonnali, A. aranica and A. aurelioi) are relatively congruent with the biogeographic sectorizations proposed up till now, although don’t fit well with the hierarchy of these subdivisions. The differentiation degree among conspecific populations is significatively correlated with their separation distances. Our sectorization from the Alpine belt developed in basis to the alpine carabofauna is the first model specifically proposed for these biota. It clusters the alpine massifs in four groups: a) Occidental Pyrenees; b) Central Pyrenees (with a central portion s. str., an aranese portion, and a preoriental portion); and c) Eastern Pyrenees.
Arribas, O.J. (1998) -
Die Pyrenäenarten von Archaeolacerta s.l. (A. aranica, A. aurelioi und A. bonnali) unterscheiden sich von den übrigen Arten der Eurasiatischen Radiationslinie (sensu MAYER & BENYR 1994) durch den Besitz knöcherner Rippen im Bereich des dritten Präsakralwirbels. Die drei Pyrenäenarten zeigen in der Morphologie von Postorbitale und Postfrontale jeweils osteologische Verschiedenheiten, die Artniveau erreichen und ihre zweifelsfreie Unterscheidung erlauben. Auch finden sich Unterscheidungsmerkmale im Schultergürtel (Klavikel und Sternalfontanella).
Apomorphe Merkmale innerhalb der kaukasischen ´Lacerta` saxicola – Gruppe stellen deutliche Unterschiede sowohl gegenüber den Archaeolacerten Europas (fast immer einzusätzlicher Präsakralwirbel) als auch gegenüber Apathya (ein zusätzlicher Präsakralwirbel und eine verminderte Anzahl von Prämaxillarzähnen) dar. ´L.` chlorogaster unterscheidet sich von den übrigen kaukasischen Formen klar durch die Zahl der Prämaxillarzähne und Form der Interklavikel. Die parthenogenetischen Taxa zeigen deutliche Ähnlichkeiten (Modalwerte der Wirbelzahl, Morphologie des Postorbitale) zu ihrem Elternarten, besonders zur weiblichen Elternart.
Aufgrund der osteologischen Befunde wird deutlich, daß die europäischen, vorläufig in der Sammelgattung Archaeolacerta s.l. vereinigten Taxa keine natürliche Einheit darstellen. Die Formen der Pyrenäen und des Kaukasus erscheinen osteologisch deutlich differenziert, während die iberischen, balkanischen und tyrrhenischen Taxa in den untersuchten Strukturen nur synplesiomorphe Merkmalsausprägungen gemeinsam haben, die keine Aussagen über ihre Verwandtschaft ermöglichen.
Auf Grundlage allen verfügbaren Materials werden neben einer umfangreiche Diagnose (Biometrie, Pholidose, Osteology, Karyotyp, Allozymdifferenzen, hemipenismorphologie) Angaben zu Verbreitung, Sexualdimorphismus und geographischer Variabilität von ´Lacerta` aurelioi ARRIBAS, 1994 vorgelegt. Pholidotische und metrische Merkmale werden dabei mit uni- und multivariatenn statistischen Methoden untersucht.
`Lacerta` aurelioi ist in einem kleinen Bereich der östlichen Zentralpyrenäen endemisch. Die Art bewohnt die spanischen Gebirgsmassive Montroig und Pica d´Estats i. w. S. (inklusive Serrat de Capifonts) sowie NW-Andorra (Gebirgsstöcke Coma Pedrosa und Tristaina), woe sie zwischen 2100 m bis fast 2940 m Höhe gefunden wird. Dort kommt die Eidechse hauptsächlich auf kambrio-ordovizischen quarzitischen Schiefern und Phylliten, aber auch auf schwach metamorphosierten Staurolitschichten und schiefrigem Cordierit und Andalusit vor.
Morphologisch zeiegen die untersuchten Populationen nur schwache Unterschiede auf infrasubspezifischem Niveau. Auch lassen sich keine Korrelationen zwischen Differenzierungsgrad und geographischer Lage oder Entfernung der Populationen feststellen: Die Subpopulationen eines gebirgsstockes (verschiedener benachbarter Hänge) sind häufig untereinander stärker verschieden als gegenüber Subpopulationen anderer Gebirgsstöcke, sodaß geographisch weiter voneinander entfernte Populationen einander morphologisch stärker ähneln können als geographisch nahe Populationen.
Ein denkbares Szenario der Artbildung, das pleistozene Überleben und die Ausbreitung sowie die gegenwärtige Verbreitung der Art werden diskutiert. Die heutige Verschiedenheit der Populationen wird auf ihre holozäne Isolation zurückgeführt. Die Südhänge des Montroig Massives werden als ein würmzeitliches Refugium angesehen, von dem aus die Art ihr gegenwärtiges Verbreitungsgebiet, in dem die verschiedenen Populationen isoliert wurden, besiedelte.
Iberolacerta bonnali is endemic from the Central Pyrenees, where inhabits from the Arriel Massif in the west, to the Aigüestortes mountains in the east. His range is characterized by the greater size of the mountain massifs which inhabits, and by to present granitic (igneous) rock substrates. By other side, his localities are also characterized by lesser insolations and sun radiations, as corresponds to the pyrenaean parts with greater atlantic influence. I. aranica is endemic from the Mauberme Massif and its spurs. His area appears characterized by greater snow accumulations both in winter and spring. I. aurelioi is endemic from the Montroig, Pica d`Estats and Coma Pedrosa massifs. His area is characterized by the greater insolations and sun radiations, and by the presence of lesser alpine belt surface and absolute heights of the inhabited massifs. Also, he inhabits only paleozoic rocks (mainly cambroordovician quarzites). In respect to other species present in the alpine belt: Lacerta agilis appears in localities characterized by high insolation and sun radiations in respect to other pyrenaean lizards. Zootoca vivipara do not present specific traits as lives in all the pyrenaean range and frequently sympatrically with all the other species here studied. Podarcis muralis appears in the comparatively more outer parts, with greater insolations, from the Pyrenees.
Arribas, O.J. (1999) -
Archaeolacerta s. l., in the currently utilized sense results to be a polyphyletic taxon, composed of three monophyletic and phylogenetically well differentiated taxa: a)-Archaeolacerta (s. str.) Mertens, 1921, which includes three species: A. bedriagae, A. oxycephala, and A. mosorensis, distributed by the northern part of the Central Mediterranean (Corsica, Sardinia, and west Balcanic Peninsula). His most related genera seem to be Teira (his adelphotaxon in our study), Apathya, and Omanosaura. b)-Iberolacerta gen. nov., with two subgenera: Iberolacerta s. str. and Pyrenesaura subgen. nov. includes six species: I. monticola, I. cyreni, I. bonnali, I. aranica, I. aurelioi, and I. horvathi. They are distributed by the mountains of Western Europe (Iberian Peninsula, Pyrenees, Central and Eastern Alps and the north of the Dinaric Chains). His adelphotaxon are the Caucasian and Near East species of Darevskia gen. nov. c)-The third taxa is Darevskia gen. nov., which includes the species of the `L.` saxicola complex besides `L.` derjugini, `L.` praticola, and `L.` chlorogaster. They are distributed by Eastern Europe, Caucasus, and Near East. Other clear relationships among some taxa of the Eurasian Radiation [= `Eurasische linie` from (Mayer and Benyr, 1994)] have been found: Algyroides seems to be the sister group of the species considered of uncertain phylogenetic relationships belonging to the `L.` danfordi-laevis group and of Podarcis. `L.` brandtii seems to be a very primitive species within the Eurasian Radiation. Omanosaura and Apathya appear as sister taxa. `L.` graeca occupies a very basal position in the sister group of Podarcis and relatives (see above). `L.` andreanskyi is the sister species of the Darevskia nov. and Iberolacerta nov. clade. The assimilation of `L.` andreanskyi to Teira is very problematic. Also the `L.` parva group seems to be related to Timon. Some groups of species like `L.` brandtii, `L.` parva-fraasii and `L.` danfordi-laevis probably merit generic rank, but it is necessary a more deep study before to take a decision.
Die Peña de Francia Gebirgseidechse (´L.` cyreni martinezricai) unterscheidet sich von anderen Formen ihrer Art (Gredos: ´L.` cyreni castiliana, Guadarrama: ´L.` cyreni cyreni) durch die signifikant verminderte Häufigkeit des Auftretens eines Schuppenkontaktes zwischen Rostrale und Internasale. Weiters zeichnen sich Männchen und Weibchen des Peña de Francia Taxons gewöhnlich durch blaue Ozellen in der Schultergegend und braune Rückenfärbung aus, während ´L.` cyreni anderer Populationen keine Ozellen tragen und sehr häufig grünrückig sind. Neben diesen sehr auffälligen (und evolutiv bedeutsamen) Farbmerkmalen finden sich signifikante aber nicht sehr starke Pholidoseunterschiede (Anzahl Rückenschuppen-Längsreihen, Bauchschuppen-Querreihen, Circumanalschilder). Das Vorkommen von ´L.` cyreni martinezricai ist auf die Gipfelregion eines isolerten Berges begrenzt. Die Populationsgröße (~ 50 adulte Individuen) nimmt ständig ab. Schutzmaßnahmen sind dringend erforderlich.
Arribas, O.J. (2000) -
Iberolacerta bonnali (LANTZ, 1927), ein Endemit der axialen Zentralpyrenäen, bewohnt die Gebirgsstöcke Arriel, Balaitous, Vignemale (oder Comachibosa), Panticosa, Monte Perdido, Punta Suelza,Posets, Maladeta, Ballibierna und die Gebirge des Sant Maurici-Aigüestortes Nationalparks (Besiberris, Muntanyó de Llacs, Peguera und Encantats) samt einer kleinen nördlichen Exklave in den Massiven von Neouvielle und Bigorre. Das Artareal umfaßt das Gebiet zwischen den Pässen von Portalé (im Westen) und Bonaigua (im Osten). Die Eidechse lebt in der alpinen Region, gewöhnlich in Höhen oberhalb 2000 m (zwischen 1700 m und 3062 m) und kommt dort in kleinen bis mittelgroßen, gewöhnlich eng begrenzten Populationen auf unterschiedlichem Gestein (Kalk, Schiefer, seltener und weniger dicht auf Gneis und Granit) und häufig in der Nähe von Seen und Bächen vor.
Die Stichproben von Posets und Maladeta erscheinen als morphologisch zentral bzw. intermediär. An sie lassen sich die übrigen Stichproben aus verschiedenen Richtungen anschließen. Unter den gut repräsentierten Stichproben scheinen die von Bigorre und Ballibierna denen von Maladeta, Monte Perdido und Posets am nächsten zu stehen. Unter den weniger gut vertretenen Samples nähern sich die von Neouvielle und Arriel an Bigorre an. Punta Suelza Exemplare sind intermediär und nahezu nicht von M. Perdido und Posets Tieren unterscheidbar. Die Stichproben Maladeta, Besibierri, Aigüestortes and Ballibierna sind alle gleichermaßen gegenüber den intermediären Populationen differenziert. Insgesamt entsteht der Eindruck, daß die U-förmige Gebirgsformation von Posets, Maladeta und Ballibierna von einer Gruppe „zentraler Populationen“ bewohnt ist, von denen sich drei Stichproben morphologisch am stärksten abheben: Bigorre im Norden, M. Perdido westlich und Aigüestortes im Osten. Dieses Modell positioniert einen möglichen Refugialstandort der Art im Würm auf die Südhänge jenes U-förmigen Gebirgskomplexes, doch bleibt die Möglichkeit anderer kleiner Refugialräume, etwas im Gebiet von Aigüestortes. Das Szenario der Ausbreitung aus diesen Refugialräumen am Ende der letzten Vereisung wird dargestellt, eine das Gesamtmaterial einbeziehende vollständige Diagnose von I. bonnali wird gegeben.
The study techniques permit us to check, from a morphological point of view, the existence of three well differentiated species in the Pyrenees: Iberolacerta aranica (Arribas, 1993), Iberolacerta aurelioi (Arribas, 1994) and Iberolacerta bonnali (Lantz 1927). ANOVA detects differences which accumulate aiming populations of different species. Characters with greater discriminative power among taxa are dorsalia, gularia, ventralia and the relative diameter of the tympanic scale. In the canonical analysis, first axis separates central-eastern pyrenaean populations (I. aurelioi) in respect to central pyrenaean ones (I. bonnali and I. aranica). Second axis separates central pyrenaean populations living in a small isolate at the northeast to the Garona river (I. aranica) from the western ones living in the main part of the pyrenean range (I. bonnali). I. aurelioi is characterized by high values ol ventral pattern, gularia, dorsalia and ventralia; and Iow values of masseteric and tympanic diameters, (temporal areas covered only by small tiny scales are not uncommon), generalyzed absence of contact among postocular and parietal scales and a lower number of circumanals and femoral pores. This species can be distinguished unequivocally also by his yellow belly, which is absent in the other pyrenaean species. I. aranica is characterized by greater diameters of the masseteric plate, relatively well developed ventral pattern. low values of dorsalia. ventralia, circumanalia and relatively short fore and hindlimbs. Usually shows a very characteristic scale disposition in temporal area, with three great scales (masseteric. tympanic and a great wedge-shaped intermediate scale). I. bonnali has intermediate pholidotic values among I. aurelioi and I. aranica. The three species show a good discrimination degree among them (superior to 75 %). The best discrimination is for I. aurelioi from which specimens rarely are mistaken with the other two species. Between I. aranica and I. bonnali there is a more imprecise discrimination, although ever superior to 75 % and more marked in males. In respect to the intraespecific variability among species from which more than one massif has been studied. I. aurelioi is the one which shows samples with the best discrimination degree among it (correct classification percentages from 71.42 to 91.6 %). In I. bonnali intracspecific discrimination is lower. Only the males from Bigorre reach the 75 % of correct classification, which suggests that this species is relatively uniform.
Arribas, O.J. (2001) -
Iberoalacerta aranica (ARRIBAS, 1993) ist ein Endemit der Zentral-Pyrenäen, der ausschließlich das Maubèrme Massiv und dessen Ausläufer zwischen Val d´Aràn (Spanien) und der Ariège (Frankreich) bewohnt. Vorkommen der Art sind in fünfundzwanzig 1 km x 1 km U.T.M. Rasterfeldern nachgewiesen; das mögliche Verbreitungsgebiet bedeckt maximal 36 derartige Rasterfelder. Die Eidechse bewohnt Schutthalden, Geröllfelder und steiniges Grasland (meist auf paläozoischen Schiefern und Kalken) im alpinen Vegetationsgürtel zwischen 1940 m (1900 m) und 2668 m Seehöhe.
Alle untersuchten Populationen mit Ausnahme deren von Orlá scheinen untereinander mehr oder weniger in Kontakt zu stehen. In Abhängigkeit vom bewohnten Gesteinsuntergrund bestehen populationstypische Unterschiede in der Rückengrundfarbe. Tiere von Maubèrme ähneln jenen von Liat sehr; beide stehen den Serra d´Armeros Populationen (Barrados und Pica Palomera) nahe. Im allgemeinen dominieren in letzteren Populationen dunkle Farbtöne, während bei Exemplaren von Maubèrme und Liuat intermediäre oder hellere Grundfarben vorherrschen. Estany de Güerri ist ein Vorkommen am äußersten Ende der Serra d´Armeros, und das einzige von dort untersuchte Exemplar scheint morphologisch den Tieren von Armenos nahezustehen.
Nur die Exemplare von Orlà unterscheiden sich deutlich von allen übrigen. Sie besitzen eine hellere Rückengrundfarbe mit deutlicher dunkler Färbung. Pholidotisch unterscheiden sie sich von den übrigen Populationen an der Körperunterseite (vor allem ihre größeren Gularia und Ventralia) sowie durch kleinere Masseterica und Analia; auch haben sie relativ längere Hinterbeine. Die Unterschiede in den Extremitätenproportionen könnten im Zusammenhang mit der gut ausgebildeten Kletterfähigkeit und dem bewohnten Gesteinstyp gesehen werden. Die Populationen von Orlà sind entweder tatsächlich isoliert oder sie stehen mit Maubèrme Tieren über einen sehr schmalen Korridor in Verbindung.
Arribas, O.J. (2002) -
Arribas, O.J. (2004) -
Iberolacerta aurelioi (ARRIBAS, 1994) ist eine in ihrem Bestand gefährdete Eidechsenart mit einer sehr kleinräumigen Verbreitung. Wesentliche Merkmale ihrer Fortpflanzungsbiologie wie etwa die hohe Lebenserwartung (von bis zu 17 Jahren, die verzögerte Geschlechtsreife (nicht vor dem 5. Kalenderjahr bzw. 4 Überwinterungen) und sehr geringe Vermehrungsrate (ein Gelege aus 2 oder 3 Eiern pro Jahr) stehen im Zusammenhang mit der kurzen Jahresaktivitätsdauer in großen Höhen. Das Datum des Erscheinens nach der Winterruhe hing von den tatsächlichen Wetterverhältnissen im jeweiligen Jahr ab und war dann für alle Altersstufen und beide Geschlechter dasselbe. Die Paarung begann bald nach dem Verlassen der Winterquartiere (zweite Maihälfte bis erste Junihälfte). Für die Paarungsstellung war der Flankenbiß des Männchens charakteristisch. Die Eientwicklung erfolgte im Verlauf des Juni und die Eiablage fand zwischen Mitte Juni und Mitte Juli statt.
Es erfolgte nur eine Eiablage pro Jahr, wobei (1) 2 oder 3 Eier gelegt wurden, welche Embryonen in fortgeschrittenen Reifestadien (Stadium 30 bis 32 nach DUFAURE et al. 1961) Bevorzugt wurden Ablageorte unter mittelgroßen bis großen flachen Steinen in leicht geneigtem Gelände. Unter solchen Steinen fanden sich zwei bis 12 frisch abglegte Eier, die von mehreren (im Durchschnitt zwei) Weibchen gemeinsam am selben Platz abgelegt worden sein mussten. Allerdings wurde solche Ablageorte häufig von Jahr zu Jahr gewechselt, sodaß nur wenige von ihnen in aufeinanderfolgenden Jahren benutzt waren. Die Veränderungen in den Abmessungen der Eier während der Inkubation werden beschrieben. Die Eizeitigung dauerte im Labor zwischen 31 bis 44 (im Mittel 35) Tage. Im Freiland traten die ersten Schlüpflinge zwischen Mitte August und Mitte September auf. Im Freiland zerstörte Fraß durch die Larve der Fliege Sarcophaga protuberans 25 % der Gelege. 64 % der im Labor erbrüteten Eier schlüpften. Der Schlupfvorgang und das frisch geschlüpfte Jungtier werden beschrieben.
Arribas, O.J. (2005) -
Arribas, O.J. (2007) -
Arribas, O.J. (2008) -
Arribas, O.J. (2009) -
Iberolacerta bonnali (LANTZ, 1927 war von 07.00 bis etwa 16.00 Uhr GMT aktiv (gewöhnlich zwischen 09.00 und 11.00 Uhr), unabhängig von Geschlecht oder Alter. Die Orographie und das jeweils herrschende Wetter schränkten die nachmittägliche Aktivität der Eidechse ein. Im Laufe des Sommers und mit abnehmender Tageslänge verschob sich der Aktivitätsgipfel zunehmend in Richtung späterer Tagesstunden. In Kalksteingebieten begann und endete die Tagesaktivität früher als in Schiefergebieten, doch unterstützte in letzteren das tigmothermische Verhalten der Eidechsen wahrscheinlich den Prozeß des Aufwärmens.
Die Hangneigung war nicht mit der Nutzungshäufigkeit durch I. bonnali korreliert. Die häufigsten und meist genutzten Mikrohabitate waren felsige oder grasbestandene Stellen. Die Eidechsen glichen Abweichungen von der mittleren Häufigkeit dieser Strukturen in allen drei Substrattypen (Kalk, Schiefer, Grandiorit/Granit) durch Bevorzugung oder Vermeidung der verfügbaren Mikrohabitate aus. In Schiefergebieten war Fels weniger häufig und wurde positiv selektioniert, während lose Gesteinbrocken relativ häufig waren und weniger angenommen wurden. Im Kalkgestein fehlte die Buschvegetation und die häufigsten Mikrohabitate, Fels und Gras, wurden etwas weniger angenommen, während umherliegende Steine selten auftraten und vermehrt als Aufenthaltsort gewählt wurden. Auf Grandiorit/Granit-Substrat wurden Grasstandorte weniger angenommen als Fels und loses Gestein.
Die Körpertemperaturen aktiver Eidechsen reichten von 20.8°C bis 35.2°C (Mittel 28.96°C ± 0.24°C) und waren etwas stärker mit der Substrattemperatur (r = 0.37) als mit der Temperatur der Umgebungsluft korreliert (r = 0.33). Signifikante Unterschiede zwischen den Geschlechtern oder Altersgruppen oder im Hinblick auf unterschiedliche Substrattypen waren nicht feststellbar. Iberolacerta bonnali waren effektive Thermoregulatoren (die Anstiege der Regressionsgeraden von 0.14 bzw. 0.21 der Körpertemperatur in bezug zu Substrat- bzw. Lufttempetratur waren stark von I verschieden), aber nicht sehr präzise bei der Einstellung ihrer Körpertemperaturen (geringe Korrelationen) und damit eher Thermokonformisten (in einem breiten Temperaturspektrum aktiv), wie auch andere Iberolacerta-Arten.
Arribas, O.J. (2010) -
Der Autor legt eine vergleichende Untersuchung über Aktivität, Mikrohabitatwahl (Hangneigung, Flächenanteil von Fels, Steinen, nacktem Boden, Gebüsch und krautiger Vegetation im Lebensraum) und temperaturabhängiges Verhalten von Iberolacerta aranica (ARRIBAS, 1993), I. aurelioi (ARRIBAS; 1994) und I. bonnali (LANTZ, 1927) vor und beschreibt die Unterschiede in den untersuchten Größen im Hinblick auf Art, Geschlecht und Alter der Eidechsen. Im Aktivitätsmuster finden sich keine Unterschiede zwischen den Arten, doch differierten sie in mehreren Faktoren der Habitatausstattung ihres Aufenthaltsortes signifikant.
Iberolacerta aurelioi wurde in steilerem Gelände als I. bonnali besonders aber als I. aranica festgestellt, doch waren die Unterschiede in der Hangneigung nich signifikant. Iberolacerta aurelioi und I. bonnali bevorzugten felsige Substrate und vermieden steinige Stellen (beide Geschlechter, alle Altersklassen) und nackte Böden (nur I. aurelioi). Iberolacerta aranica bevorzugte weniger felsigen als eher steinigen un d unbewachsenen Bodengrund. Büsche waren im Habitat von I. bonnali häufiger als in jenem von I. aurelioi und I. aranica. Krautige Pflanzen (Ganz allgemein alle Formen von Vegetation) fanden sich häufiger im Lebensraum von I. bonnali als an Stellen, wo I. aurelioi oder I. aranica vorkamen.
Die drei Arten unterschieden sich voneinander in der Körpertemperatur (BT) und erwiesen sich als gute aber nicht sehr präzise Thermoregulatoren (besonders gegenüber den Subastrattemperaturen) sowie mäßige Thermokonformisten. Die Körpertemperatur war bei Iberolacerta aurelioi niedriger als bei I. bonnali (p < 0.05) und besonders I. aranica (p < 0.01). Die mittlere Körpertemperatur betrug bei I. bonnali 29.20 ± 0.30°C (22.8 – 35.2°C; n = 64), bei I. aranica 29.49 ± 0.20°C (22 – 36.5°C; n = 143) und bei I. aurelioi 28.13 ± 0.32°C (16.6 – 34.4°C; n = 109). Die Erwärmung schien bei I. aurelioi und I. bonnali hauptsächlich heliothermisch, bei I. aranica tigmothermisch zu erfolgen.
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.
Arribas, O.J. (2012) -
In this paper we deal on the ultraviolet color (invisible to us): where we can find it, the capability of animals to see it and the advantages that this color perception offers to them. As the simplest way to detect it is the photography, we describe and review how to photograph the UV, as a result of 15 years of amateur experience, searching
and testing nearly in complete blindness due to the lack of practical information about “how to do it”. We describe the different kinds of photography (chemical and digital); the cameras and objectives suitable (both astronomically expensive ones and cheap options); what are the best characteristics that the objectives should have for this purpose; the films suitable for their use in chemical photography; the different filters (current or discontinued) manufactured along the years; and the subtle combinations among the different materials to obtain pure UV photographs. This
kind of scientific photography is mainly used in forensics, forgery detection, art dermatology and less in Natural History, despite the fact that a great part of animals see this color and use it in important questions of their biology as the social behavior, mate choice or the food search.
Arribas, O.J. (2013) -
Zur Naturgeschichte von Iberolacerta martinezricai (ARRIBAS, 1996), der seltensten Reptilienart Kointinentaleuropas, liegen nur wenige Angaben vor. Das zahlenmäßige Geschlechterverhältnis betrag nach Informationen aus der Datenbank des Autors 0,74:1 (Männchen: Weibchen), während im deutlich größeren datensatz der feldbeobachtungen (dieser schließt allerdings Beobachtungswiederholungen ein) die Männchen überwogen (1,27:1).
Alla Altersgruppen und Geschlechter zeigten ähnliche, die Morgenstunden betonende Tagsaktivitätszyklen. Nachmittagsaktivität erfolgte gehäuft an schattingen Orten, wobei Sonnenexposition bei hohen Lufttemperaturen vermiden wurde. Zwei Häutungsperioden (eine im April, eine um Ende Jun und Juli) wrden beobachtet. Der Lebensraum war mäßig neiegtes, ausnahmslos von Felstrümmern und großen Blöcken, mit reichlich Moos- und Flechtenbewuchs dominiertes Gelände, was auf erhöhte Feuchte und geologische Hang-Stabilität hinwies. Besondere Vorlieben fürf einzelne Mikrohabitate waren nicht feststellbar.
Die Körpertemperatur aktiver Individuen war die höchste innerhalb der Gattung Iberolacerta: 33.31±0.12°C (Spannweite 22.8-39.2°C) ohne signifikanten Geschlechtsunterschied. Im Mittel hatten blaufärbige Männchen eine höhere Körpertemperatur als grüne (33.73±0.19°C gegenüber 32.8±0.28°C). Die Aufenthaltsorte adulter Weibchen wiesen eine höhere Lufttemperatur auf (23.77±0.20°C, Spannweite 13.8-37.1°C) als die der Jungtiere. Dioe Substrattemperaturen (30.08±0.31°C, Spannweite 16.5-52.1°C) waren an den Aufenthaltsorten derWeibchen höhetr als an dene der Männchen.
Iberolacerta martinezricai war mäßig thermokonformistisch (Spannweite der Aktivitätstemperatur: 16.4°C) und ein effektiver und ziemlich präziser Thermoregulator was die Substrattemperatur betrifft(r = 0.36, Anstieg 0.14), aber zeigte kein thermoregulatorisches Verhalten hinsichtlich der Lufttemperatur (r = 0.15, Anstieg = 0.09). Tigmothermie, Aktivität im Schatten bei hoher Lufttemperatur und Aestivation werden diskutiert-
Arribas, O.J. (2014) -
Sex dimorphism: Males and females of Iberolacerta martinezricai differ at p < 0.001 in Ventralia (MF), Forelimbs length (M>F), Hindlimb length (M>F), Pileus length (M>F) and at p<0.05 in Collaria (M>F) and Dorsalia (M>F). Batuecan lizards are overall less di-morphic than the three species of Pyrenean rock lizards.
Morphological variability: Variability Index calculated (ΣCV) for I. martinezricai was 6.8 in males and 6.0 in females. Results are very similar (although slightly smaller) to the obtained in the Pyrenean Iberolacerta spp. From two chosen characters, variation coefficient in Gularia falls also between the normal values for the bisexual species and it is a bit greater than in parthenogenetic species; or it is clearly greater as in Dorsalia. Nothing suggests a drastic reduction of the variability in the Batuecan lizard but some characters recall for a relatively small morphologic variability in some characters, in parallel with the lack of mitochondrial variability, perhaps due to its reduced effectives.
Allometry: Both in males and females, the most representative character of growth is the body length (SVL). Most other characters match very closely to this SVL with very similar coefficients: limbs, Parietal, Pileus length and the Anal plate width. Masseteric, Tympanic and Anal length also grow, but not in parallel with SVL. Anal length grows considerably and independently of body size. In the case of fe-males, body limbs have a negative allometry respect to female’s body length and Masseteric, and especially Tympanic plates have a negative allometry respect to the total growth.
Parallel variation in escalation: In males the most related characters of escalation are the two rows of Femoralia, and two characters that have contrary signs: Supraciliar Granula (but not very marked, as coefficients are very small) and specially Circumanalia (that varies with total independence). Also there is a small relationship of Supraciliar Granula with Gularia and even more distant with 4th-toe Lamellae and Circumanalia. In females on one side (first axis), both Supraciliar Granula rows are related to Femoral pores, 4th toe Lamellae, Ventralia and Femoralia; and on the other side (second axis) both sided Femoralia are lonely related with Dorsalia, Collaria and Gularia, and just inversely related to Supraciliar Granula.
Sexual maturation is probably reached when lizards are three years old (in his 4CY). Our smallest gravid female was 56.7 mm, and was probably reproducing in the spring of their 5 CY (when 4 years old). Al-though it is possible that some specimens (especially males, but also some females) reach maturity during 3CY, as it occurs in I. monticola. Growth rate (K) in the first years is 0.145696 ± 0.0633135 (95% confidence). Calculated length at infinite (maximum size if the same rate or growth curve was ap-plied throughout the lifetime of the animal) would be: L∞ = 70.036 ± 13.7598 (95%). This estimate of the asymptotic size (70.036 mm) is very close to the true maximum size found in the total sample of adults (69.31 mm). Growth decreases progressively and the resulting curve is fairly straight, being the growth not far from linear and very similar in both sexes.
Longevity: The biggest male (SVL 68.15 mm) and the biggest female conserved (SVL 68.86 mm) had 7 growth rings (two big and five narrow) in the femur. The two wide rings correspond to the two first years of rapid growth (2CY and 3CY), but when lizards reach sex maturation growth decreases, being latter rings very narrow and even difficult to count. Both specimens had presumably 8 years old (they are 9CY animals: the birth year and 7 growths seasons, and captured just in the beginning of their 9 CY).
Tail autotomy: Iberolacerta martinezricai hatchlings are able to break their tails from birth. Observed hatchlings (1CY) in nature had intact tails. In 2CY, 3CY and 4CY already appear broken and regenerated tails in some specimens. In adults, there are less than a quarter of specimens with apparently unbroken tails, and the others fall in the different categories of regenerated ones (tail tip, middle of the tail or the whole tail from basis) in proportions similar to one quarter. There are no differences between both sexes in the different categories of broken tails, but there are significant differences between juveniles and adults. Females have a slightly big percentage of broken tails from its basis. Bright colored tail in juveniles (bluish in I. martinezricai), is lost during their 2CY. It seems to be an antipredatory strategy. Hatchlings of the Batuecan lizard are able to attract attention on its tail by a striking whirl of its distal part, unlike adults that are unable with their stiffer tail (which can move only in very limited cases of danger or excitement).
Tail regeneration in 706 Iberolacerta (Pyrenesaura): 300 I. bonnali, 212 I. aranica and 194 I. aurelioi was studied, distinguishing four situations: 0) intact tail; 1) distal part (point) of the tail regenerated; 2) Tail regenerated from its half part and 3) tail broken and regenerated from its basis. The incidence of autotomy (tail break) in Iberolacerta (Pyrenesaura) runs in adults from a 47.5% up to a 68.3%. In juveniles is sensibly lower and goes from 10.9% to 23.8%.In I. bonnali percentage of autotomized tails is almost identical in both sexes (48% in males and 47.5% in females), whereas in juveniles is only 22.4%, without differences between adults but yes between adults and juveniles. In I. aranica, percentage of autotomized tails is 68.3% in males (fairly high), a 55.3% in females and a 10.9% in immatures. There are not significant differences between males and females (despite to be near to significance) but exist between adults and juveniles. In I. aurelioi males have a break percentage of 54.4%, females 59.8% and juveniles 23.8%, without differences between males and females, and contrary to the other two Pyrenean species, also inexistent between adults and juveniles. Comparing the three species among them, only males differ (the difference arises between I. bonnali and I. aranica), and all the other comparisons among females and juveniles are not significant. Juveniles of I. aurelioi (only any population of I. aranica or rarely and few vivid in I. bonnali) present bluish of greenish tails that do not broke until some days after birth and are able to wave sinuously in order to attract the predators attention upon it. Also, the known and possible predators are reviewed.
Arribas, O.J. (2017) -
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.
Arribas, O.J. (2018) -
Der Autor untersuchte das Fortpflanzungsgeschehen von Iberolacerta martinezricai (ARRIBAS, 1996). Kopulationen begannen Ende April. Im Juli trugen alle Weibchen zahlreiche frische Paarungsnarben auf Bauch und Schwanzbasis. Iberolacerta martinezricai ist monoestrisch. Die Eiablage erfolge im Juli und Anfang August, am häufigsten gegen Ende Juli. Das Eiablageverhalten wird beschrieben. Eier wurden zwischen 9:00 und 15.30 Uhr (GMT) gelegt, wobei der ganze Ablagevorgang 33-42 Minuten (Durchschnitt: 36:57 Minuten) dauerte. Die Kopf-Rumpf-Länge reproduktiver Weibchen betrug 56.7 – 69.71 mm. Die Gelege umfaßten zwei bis sechs Eier (Median und Modalwert: 4, Mittel: 4,23). Die abgelegten Eier klebten nicht aneinander. Biometrische Daten von 49 Eiern (darunter 12 Laboraufzuchten) werden angegeben. Während des Großteils der Inkubationsphase nahmen Eivolumen und Eimasse durch Wasseraufnahme kontinuierlich zu. Zu Begin der Inkubation erfolgte ein rascher Anstieg des Ei-Querdurchmessers (dieser wurde während der Ablage durch die Enge des Geburtskanals verringert). Die Inkubation dauerte 33-42Tage (im Mittel 37.75 Tage). Das Schlüpfen erfolgte zwischen 3:00 und 22:00 Uhr (GMT) (im Mittel 9.40 Uhr) und dauerte 4.10 – 12 Stunden (im Mittel 6.9 Stunden). Die Merkmale des Schlüpflings werden beschrieben. Der Eizahn bestand für acht bis 36 Stunden (im Mittel 23.6 Stunden nach Schlupfbeginn. Schlüpflinge können von Geburt an mit ihrem leuchtend blauen Schwanz wedeln und diesen autotomieren. Diese auffällige Schwanzfärbung verschwand im Laufe des zweiten Kalender-Lebensjahres.
Arribas, O.J. (2019) -
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.
Arribas, O.J. & Ananjeva, N.B. & Carranza, S: & Doronin, I.V. & Harris, D.J. & Orlov, N.L. & Orlova, V.F. (2017) -
Busack et al. (2016) concluded that the name Darevskia should be attributed to Arribas (1999) instead
of Arribas (1997), rendering the latter name a junior synonym of Caucasilacerta Harris et al.
(1998). In this paper, we provide a detailed answer to Busack et al. (2016) and argue that Arribas
(1997) fulfilled the publication requirements of the International Code of Zoological Nomenclature
(ICZN) in place in 1997 and, therefore, Darevskia Arribas, 1997 is a valid name. In order to clarify
this issue, a case has been submitted to the ICZN. To avoid unnecessary confusion in Old World
Lacertid taxonomy, we recommend that until the commission rules on this case, Arribas (1997)
should continue to be considered the authority of both Darevskia and Iberolacerta.
Arribas, O.J. & Carbonero, J. & Lizana, M. (2008) -
Arribas, O.J. & Carranza, S. (2004) -
Iberolacerta cyreni martinezricai is elevated to the species level (I. martinezricai) based on both
morphological and molecular data. The phylogenetic analysis using two mitochondrial and one
nuclear gene shows I. martinezricai is more closely related to I. monticola than to I. cyreni. A multivariate
analysis of the morphological data also supports the affinities between I. martinezricai and
I. monticola but, at the same time, clearly indicates that I. martinezricai is morphologically distinct
from both I. monticola and I. cyreni. The molecular data suggests I. cyreni and the clade formed by
I. monticola + I. martinezricai split approximately 6.1 Mya, during the Mesinian Salinity Crisis,
when climatic conditions around the Mediterranean area changed dramatically as a result of the
desiccation of the Mediterranean Sea. Separation between I. martinezricai and I. monticola
occurred approximately 2 Mya but, with at least two equally plausible alternative hypotheses, their
biogeography is still unclear. New data on the habitat and distribution of I. martinezricai indicates
its distribution area is very small (12–15 km2), and that it lives in a climatically extreme habitat for
this kind of mountain species. As a result of that and the low numbers of individuals, I. martinezricai
is considered here as Critically Endangered.
Arribas, O.J. & Carranza, S. (2009) -
Arribas, O.J. & Carranza, S. (2012) -
Arribas, O.J. & Carranza, S. & Odierna, G. (2006) -
A new species of Iberolacerta is described from the Montes de León (northwest Iberia). This new
species, Iberolacerta galani sp. nov., is characterized by its relatively large size, high number of
blue ocelli on the shoulders and the relatively frequent contact or near-contact between the
supranasal and the first loreal scale, the fairly straight squamosal bone (only curved on its posterior
part), a unique karyotype in Iberolacerta combining 2n=36 chromosomes, an L-type NOR and
differentiated W and Z sex chromosomes, and unique mitochondrial DNA sequences for the
cytochrome b and 12S rRNA genes.
The correlation analyses show that morphology in general, but especially scalation, is strongly
correlated with the amount of precipitation during the months of lizard activity, which suggests that
these are not good taxonomic characters, and that other characters apparently independent of the
climate like for instance osteological, karyological and DNA features are much more reliable in
delimiting species boundaries in Iberolacerta.
According to our phylogenetic analyses, I. galani nov. is part of a very well supported clade
that originated around 2.5 mya and also includes I. monticola and I. martinezricai. Phylogeny
suggests I.martinezricai might be the sister taxon to I. galani nov. from which it split approximately
2 mya, at the beginning of the Pleistocene. The clade containing I. galani nov., I. martinezricai and
I. monticola was probably widely distributed across western Iberia during moderately cool and
moist phases of the Pleistocene, but it was probably restricted to its present range as a result of the
general temperature increase during the Holocene and competition with other lacertid lizards.
Iberolacerta galani nov. is endemic to the Montes de León, where it is isolated from the other
species of the “monticola-group” by the Duero and Miño-Sil Rivers, but particularly by the Bibei
Arribas, O.J. & Galan P. & Remon N. & Naveira H. (2014) -
Iberolacerta populations from the Northern Montes de León (NML) were studied by means of external morphology (scalation and biometry), osteology and genetics (mtDNA and microsatellites), searching for their homogeneity (“intrazonal analysis”) and, once verified, comparing them with Iberolacerta monticola s. str. (from Central Cantabrian Mountains) and I. galani (from Southern Montes de León) (“extrazonal analysis”) from neighboring areas. Our “intrazonal analysis” revealed discordances between the different approaches, especially the patterns of variation of nuclear microsatellites (congruent with external morphology) and mtDNA, namely a very low nuclear differentiation between relatively highly differentiated mtDNA lineages. The morphological approach was unable to discriminate any of the populations as significantly different from the others in the NML. Mitochondrial DNA revealed a haplotype lineage closely related to I. galani (MNL-II in our text) in some specimens of Sierra de Villabandín and Suspirón, but these populations are morphologically indistinguishable from the main part of the other populations that belong to lineage NML-I, phylogenetically closer to I. monticola. After a separation from I. monticola ca. 1.8 Mya, the populations in this geographic region must have suffered at least two different waves of gene flow from I. galani, the second one not much later than 0.5 Mya. Microsatellite results indicate that all the NML populations are genetically similar in terms of their nuclear genomes, independently of their mitochondrial differentiation (NML-I vs. NML-II haplotype groups). Since all the morphological and microsatellite evidences point towards the fact that, independently of the mitochondrial haplotypes that they bear (NML-I or NML-II), there is only one taxon in the area, we describe it as: Iberolacerta monticola astur ssp. nov. Concerning the relationships of I. m. astur ssp. nov. with I. monticola s. str. and I. galani (“extrazonal analysis”), in the female analyses the new taxon centroid is closer to I. monticola s. str. than to I. galani (more similarity with I. monticola s. str.), whereas in the male analyses the relationship is just the contrary (closer to I. galani, paralleling the direction of the hypothesized past hybridization). Moreover, in both sexes’ ANOVA, I. m. astur ssp. nov. results more similar (less P<0.05 differences) to I. galani than to I. monticola s. str. Osteologically, I. m. astur ssp. nov. is slightly more similar to I. monticola s. str. than to I. galani, especially in the squamosal bone, which is regularly arched (primitive shape). Genetically, as indicated above, the NML populations can be subdivided in two groups according to their mitochondrial DNA, namely NML-I (bearing clearly differentiated haplotypes, phylogenetically closer to I. monticola) and NML-II (whose haplotypes could have been mistaken for those of an I. galani population). This mitochondrial subdivision has at most a subtle nuclear correlate, however. According to the nuclear microsatellite markers, all the NML populations belong to a single group (I. m. astur ssp. nov.), which would be more similar to I. galani than to I monticola, with NML-II populations lying closer to I. galani than those from the NML-I group and, correspondingly, more distant from I. monticola. The discordant phylogenetic signal of mitochondrial and nuclear markers is discussed in terms of past introgression events and sex-biases in phylopatry and dispersion in these species. Iberolacerta monticola astur ssp. nov., inhabits the Northern Montes de León (Sierra de Gistreo sensu latissimo): Gistredo, Catoute, Tambarón, Nevadín, Villabandín (or Macizo del Alto de la Cañada), Arcos del Agua (or Fernán Pérez), Tiendas and Suspirón, mainly in quartzite and slate rock substrates. Its current distribution, cornered in the NW of the Northern part of the Montes de León, suggests a possible competitive exclusion between this taxon and I. galani, as the galani haplotypes (NML-II) appear cornered in the most harsh and continental areas, speaking also about a, even in the past, very limited presence of this species in the area that probably was soon absorbed by I. m. astur ssp. nov. (with NML-Concluding, it seems that the current main distribution area of I. m. astur ssp. nov. (especially the typical NML-I) gravitates around what was the divisory between watersheds in the past, later shifted to the East during the Quaternary. Eastern known limits of I. m. astur ssp. nov. do not pass away from Collado de Campo Lamoso (1500 m), which today is perfectly suitable for the species, but during the Pliocene and the main part of the Pleistocene, constituted a barrier across which the two northern immediate valleys drained to the southern slopes. The West-East continuity of this massif during the end of the Miocene was broken by changes in the drainage across this pass in the Pliocene (geological datation uncertain). Although nowadays the pass to the East (to the Filera Massif, 1879 m) is possible for Iberolacerta, the prospections in these drier limestone areas had been unfruitful. In the north of these Sierras, the species can reach up to Cascaros peak (1854 m), but this extreme has to be confirmed.
Arribas, O.J. & Galán, P. (2005) -
The three lacertid lizards species of the Iberolacerta genus, which have recently been described or recognised as different species, are reptiles that live strictly at the highest altitudes in Europe, from elevations of near 1900 up to more than 3000 m a.s.l. in the Pyrenees (Spain, Andorra, France). In this paper, the reproductive cycle and reproductive characteristics of these species are described for the first time. Data were obtained from field studies carried out in different high-mountain Pyrenean locations from 1989–2002. In addition we also conducted hatching studies in laboratory. Due to the harsh climatic conditions in the high mountains the activity cycle of these species is very short, just over 4 months, from mid-May to late September or beginning of October. This affects their reproductive cycle (i.e., only one annual egg-clutch is produced), as well as other reproductive characteristics, as the existence of a very advanced embryonic development at oviposition, which is interpreted as an advanced stage in the tendency towards viviparity, and diverse life history characteristics as a very little annual growth which greatly delays sexual maturity to 4 years in males and 4–5 years in females. Clutch size correlates significantly with female snout-vent length (SVL) in all three species. The average clutch size is 2.53 eggs in I. aurelioi (the smallest species), 3.03 in I. bonnali and 3.44 in I. aranica (the biggest species). Notwithstanding the differences in egg number the three species have a similar egg volume. The incubation period in the laboratory is very short with an average of 30–36 days among the species. The low reproductive potential observed in these three species and revealed in this paper is a strongly threatening factor which, together with their extremely reduced distribution area, endangers these endemic, rare and very threatened species.
Arribas, O.J. & Martinez-Rica, J.P. (1997) -
Arribas, O.J. & Odierna, G. (2004) -
Iberolacerta (cyreni) martinezricai has a karyotype of 2n = 36, all acrocentric macrochromosomes. Nucleolar Organizer Regions (NORs) are interstitially situated on a medium small (MS-type) chromosome, a derived character shared with I. cyreni, which differs in details of W-sex chromosome: W is heteromorphic and heterochromatic in I. cyreni, while it is homomorphic and euchromatic in I. (c.) martinezricai. Osteologically, it is characterized by the presence of a triangular-shaped expansion of the squamosal towards the supratemporal fenestra, and by the presence of seven (instead of six) posterior short ribbed presacral vertebrae. These odd characters probably became fixed in this relic taxon by imbreeding. Its differences and affinities are discussed, proposing a specific status for this taxon.
Ayllón Lopez, E. & Barbera Luna, J.C. (2011) -
Baeckens, S. & Edwards, S. & Huyghe, K. & Van Damme, R. (2015) -
Animals communicate via a variety of sensory channels and signals. Studies on acoustic and visual communication systems suggest that differences in the physical environment contribute to the variety of signalling behaviour, with species investing in those signals that are transmitted best under the local conditions. Whether or not environmental tuning also occurs in chemical communication systems has received much less attention. In the present study, we examined the effect of several aspects of the physical environment on the chemical communication system of lacertid lizards (family Lacertidae). The numbers of femoral pores are used as a proxy reflecting how much a particular species invests in and relies upon chemical signalling. Femoral pores are specialized epidermal structures that function as a secretion channel for the waxy substance produced by glands. In some lacertid species, the secretion carries infochemicals that play an important role in social communication. The number of femoral pores varies considerably among species. We have compiled data on femoral pore numbers for 162 species and tested for the effects of climate and substrate use. After correcting for body size and taking the phylogenetic relationships among the species into account, we found no effect of climate conditions or latitude on species pore numbers. Substrate use did affect pore numbers: shrub-climbing species tended to have fewer femoral pores than species inhabiting other substrates.
Baeckens, S. & Garcia-Roa, R. & Martin, J. & Damme, J. van (2017) -
Lizards communicate with others via chemical signals, the composition of which may vary among species. Although the selective pressures and constraints affecting chemical signal diversity at the species level remain poorly understood, the possible role of diet has been largely neglected. The chemical signals of many lizards originate from the femoral glands that exude a mixture of semiochemicals, and may be used in a variety of contexts. We analyzed the lipophilic fraction of the glandular secretions of 45 species of lacertid lizard species by gas chromatography/mass spectrometry. The proportions of nine major chemical classes (alcohols, aldehydes, fatty acids, furanones, ketones, steroids, terpenoids, tocopherols and waxy esters), the relative contributions of these different classes (‘chemical diversity’), and the total number of different lipophilic compounds (‘chemical richness’) varied greatly among species.We examined whether interspecific differences in these chemical variables could be coupled to interspecific variation in diet using data from the literature. In addition, we compared chemical signal composition among species that almost never, occasionally, or often eat plant material. We found little support for the hypothesis that the chemical profile of a given species’ secretion depends on the type of food consumed. Diet breadth did not correlate with chemical diversity or richness. The amount of plants or ants consumed did not affect the relative contribution of any of the nine major chemical classes to the secretion. Chemical diversity did not differ among lizards with different levels of plant consumption; however, chemical richness was low in species with an exclusive arthropod diet, suggesting that incorporating plants in the diet enables lizards to increase the number of compounds allocated to secretions, likely because a (partly) herbivorous diet allows them to include compounds of plant origin that are unavailable in animal prey. Still, overall, diet appears a relatively poor predictor of interspecific differences in the broad chemical profiles of secretions of lacertid lizards.
Balletto, E- (2005) -
Bank, J., Kruyntjens, B. & P. Paulissen (1982) -
The authors give a description of the herpetofauna of some area´s in Yougoslavia they visited in june 1979. Some attention is given to the habitat of several species. The differences between certain island and mainland populations of some Lacerta- and Podarcis-species are discussed to some extend.
Barbadillo Escrivá, L.J. (1985) -
Two populations of Lacerta monticola cyreni from the sierra de Gredos (Avila, central Spain) have been studies. Eight egg-layings in captivity and one from natural habitat have been analyzed. Nineteen specimens finally hatched with represents a reproductive success of about th 40 %. Egg-laying occurs in July and the clutches range from 3 to 9 eggs, with an average of 5,4 eggs per clutch. Hatching begins around the last days of August and last until the end of September. The newborn females are larger in S.V.L. (snout-vent length) than males. The colour and pattern of the hatchlings is very similar within the same clutch, but these characteristics differ from one clutch to another. The degree of calcification of the eggshells along with the environmental humidity during the period of incubation have proved to be two important factors to take into consideration in order to obtein a high reproductive success.
Barbadillo, L.J. & Lacomba, J.I. & Pérez-Mellado, V. & Sancho, V. & López-Jurado, L.F. (1999) -
Barbadillo, L.J. Valdemorgo, D.G. & Sánchez-Herraiz, M.J. (1997) -
Barthe, L. (Coord.) (2014) -
Bas-López, S. (1980) -
Bas-López, S. (1982) -
The present work studies the herpetological community of Caural (NW, of Spain): biogeography and ecology.
Biogeography altitudinal distribution of species, altitudinal variation in abundance of species and stratification reflect a decrease with the altitude bound to climatic reasons and reproductivity in reptiles and to lesser heterogeneity of biotopes in amphibians. There is a greater altitudinal stratification of the subcommunity of amphibians than of reptiles.
Ecology: the different physiological requirements of amphibians from land habitats and lizards are reflected in a temporal alternation (seasonal and daily) in the exploitation of the same trophical resource. The snakes occupy anoth trophical level (predators and superpredators) and the tritons live inanother habitat (subaquatic) with different resources and they have a different diet.
The problem of interspecific competition is dealt with in two concrete cases: Lacertidae and Triturus. Great differences in size or corporal shape and behaviour permit coexistence but in the opposite case they separate themselves throughout the area, distributing the biotopes according to their physiological preferences.
There are cases (Lacerta schreiberi and Triturus boscai) in which there appears to be a spatial displacement due to a diffuse competition with congenital species.
One highlights the importance of the habitat in the food diet of some species, which becomes very clear comparing the tritons with the lizards and amphibians from land habitats.
One questions the problem of stability and heterogeneity (climatic and spatial), competition and predation as factors which impel and influence the evolution of communities and species.
In relation with studies realized in other areas we see that the herpetological community of Caurel represents the transition from the Mediterranean region to the European shore region, very much in accordance with the climatic and botanical characteristics of the zone, where the seasonality plays an important role just as the glaciarism appears to have determined a reptile community evolutionally young.
Battisti, C. & Luiselli, L. (2011) -
In connectivity conservation and ecological network planning, the selection of focal fragmentationsensitive species represents a priority step. Nevertheless, despite their strategic role, selection of focal species has traditionally been carried out using charismatic and/or non objective approaches. In this way, actions of planning and conservation could be ineffective. Using as a case study Italian reptiles, we apply an expert-based approach for the selection of focal species on the basis of sensitivity to components of habitat fragmentation (habitat area reduction, increase of habitat isolation, increase of edge effect and landscape matrix disturbance) and of intrinsic ecological traits of the species (trophic level, dispersal ability, body size, niche breadth, rarity). The threshold values for each component of fragmentation defined a set of 21 focal species that can be divided into the three macro-components of human-induced habitat fragmentation (HIHF) towards which they show a sensitivity, the suitable spatial scale of populations and relative suitable habitat categories. Among these species, seven can be sampled easily with standard, low-cost field protocols. The selected species largely coincide with the species known in literature as fragmentation-sensitive.
Bauwens, D. & Garland, T. & Castilla, A.M. & Van Damme, R. (1995) -
Oganismal performance abilities occupy a central position in phenotypic evolution; they are determined by suites of interacting lower-level traits (e.g., morphology and physiology) and they are a primary focus of natural selection. The mechanisms by which higher levels of organismal performance are achieved during evolution are therefore fundamentally important for understanding correlated evolution in general and coadaptation in particular. Here we address correlated evolution of morphological, physiological, and behavioral characteristics that influence interspecific variation in sprint speed in a clade of lacertid lizards. Phylogenetic analyses using independent contrasts indicate that the evolution of high maximum sprinting abilities (measured on a photocell-timed racetrack) has occurred via the evolution of (1) longer hind limbs relative to body size, and (2) a higher physiologically optimum temperature for sprinting. For ectotherms, which experience variable body temperatures while active, sprinting abilities in nature depend on both maximum capacities and relative performance levels (i.e., percent of maximum) that can be attained. With respect to temperature effects, relative performance levels are determined by the interaction between thermal physiology and thermoregulatory behavior. Among the 13 species or subspecies of lizards in the present study, differences in the optimal temperature for sprinting (body temperature at which lizards run fastest) closely matched interspecific variation in median preferred body temperature (measured in a laboratory photothermal gradient), in- dicating correlated evolution of thermal physiology and thermal preferences. variability of the preferred body tem- peratures maintained by each species is, across species, negatively correlated with the thermal-performance breadth (range of body temperatures over which lizards can run relatively fast). This pattern leads to interspecific differences in the levels of relative sprint speed that lizards are predicted to attain while active at their preferred temperatures. The highest levels of predicted relative performance are achieved by species that combine a narrow, precise distribution of preferred temperatures with the ability to sprint at near-maximum speeds over a wide range of body temperatures. The observed among-species differences in predicted relative speed were positively correlated with the interspecific variation in maximum sprinting capacities. Thus, species that attain the highest maximum speeds are (1) also able to run at near-maximum levels over a wide range of temperatures and (2) also maintain body temperatures within a narrow zone near the optimal temperature for sprinting. The observed pattern of correlated evolution therefore has involved traits at distinct levels of biological organization, that is, morphology, physiology, and behavior; and trade- offs are not evident. We hypothesize that this particular trait combination has evolved in response to coadaptational selection pressures. We also discuss our results in the context of possible evolutionary responses to global climatic change.
Beade Toubes, E. (2019) -
The low-altitude populations of the threatenedlacertid lizard Iberolacerta monticola in A Coruña province have suffered population declines in the last decades. The present study has been done on the main population of this species in the Mandeo river basin, located in the river reserve of Chelo. Population size and density have been estimated, as well as other population parameters. We used the MARK program, which allows to use different variants of the Jolly-Seber method. In addition, we did an analysis of the sex-ratio and the structure of the population using size classes.The results indicated a high density, but limited to a reduced area with favourable habitat, surrounded by inadequate zones (forests with dense tree coverage),as well asa high mortality of immature individualsand an unequal proportion betweenimmaturesand adults. It seems to point out that this population shows insular characteristics. Other relevant data forthe survival of this population were obtained, as the apparent lack of predators, based on the low percentage of individuals with tail loss, or the reduced presence of individuals affected by ectoparasites of the genus Ixodes. The low number of observed movements points out that the individuals of this population are essentially sedentary. Some consequences for the survival of this population are set and some measures are proposed for its conservation.
Beck, P. (1943) -
Bellese, A. (2013) -
Bensettiti, F. & Gaudillat, V. (2002) -
Berroneau, M. (2010) -
Bertók, P. (2014) -
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.
Biró, G. (2017) -
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.
A viselkedésökológiai kutatásokban csak az utóbbi 15 évben ismerték fel, hogy az egyedek közötti konzisztens viselkedésbeli különbség (más szóval állati személyiség) biológiailag releváns és érvényes jelenség. Az állati személyiség léte nem magától értetődő, mivel számos módon csökkentheti a rátermettséget. Például az agresszív egyedek agresszívek maradnak olyan kontextusokban vagy szituációkban, amikor a viselkedés maladaptív, például a párjukkal és utódaikkal szemben, vagy ragadozó jelenlétében. Ezért az állati személyiség kialakulásában és evolúciójában szerepet játszó mechanizmusok azonosítása fontos célunk.
Bischoff, W. (1984) -
Bischoff, W. (1991) -
Bischoff, W. (2006) -
Blasco, M, (1974) -
Bloor, P. (2006) -
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.
Bolkay, S.J. (1921) -
Bolkay, S.J. (1923) -
Bolkay, S.J. (1924) -
Bolkay S.J. (1924) -
Bonato, L. (2011) -
Borg, J.P. ter (2004) -
Reptiles and amphibians in Portugal
The author spend two weeks in July 2002 in Beira, Portugal. The following observations were made:
Vale de Asna (Serra do Açor): the sound of several Alytes obstreticans boscai (probably), several Tarentola mauritanica, one Podarcis hispanica type 1, many Psammodromus algirus, several Chalcides chalcides striatus, larvae of Rana iberica, Triturus marmoratus marmoratus, Triturus helveticus sequeirai and Triturus boscai, one adult Rana perezi, one dead Bufo bufo, three Natrix maura (half-grown and adult).
Waterfall of Sobral (Serra do Açor): many Psam-modromus algirus, several larvae, juvenile and adult Rana iberica, many juvenile Bufo bufo.
Pena (Serra da Lousã): several Psammodromus algirus.
Mata da Margaraça (Serra do Açor): several Psam-modromus algirus, one Podarcis hispanica (probably).
Fraga de Pena (Serra do Açor): several larvae and two adult Rana iberica, one Natrix maura (probably), several larvae and three adult Chioglossa lusitanica.
Serra da Estrela: Near the top of mount Torre (between 1750 and 1993 m): many Iberolacerta monticola monticola, many larvae of Bufo species, many adult Rana perezi, one adult Rana iberica, one adult Hyla arborea molleri. Around 1200 m, one adult and one juvenile Timon lepidus, one adult Lacerta schreiberi. At 1600 m one juvenile Lacerta schreiberi.
Between Piódão and Foz d’Égua (Serra do Açor): one half-grown Natrix natrix astreptophora, one juvenile Natrix maura (probably newly hatched), several Psammodromus algirus and one adult Malpolon monspessulanus.
It is stated that Portugal is a very interesting place for spotting reptiles and amphibians in nature.
Börner, A.-R. (1975) -
Bosch, H.A.J. in den (1994) -
Boulenger, G.A. (1905) -
Boulenger, G.A. (1910) -
Boulenger, G.A. (1913) -
Bovero, S. & Canalis, L. & Crosetto, S. (2013) -
Brana, F. (1984) -
Braña, F. (1996) -
Die Arbeit basiert auf der Messung verschiedener metrischer Charaktere (Kopf-Rumpf-Länge, Kopflänge, maximale Kopfhöhe, maximale Kopfbreite und Rumpf-Länge) an insgesamt knapp 500 nordspanischen Eidechsen. In die Analyse gingen Mesungen an Podarcis bocagei, Podarcis hispanica, Podarcis muralis, Lacerta monticola, Lacerta vivipara, Lacerta lepida, Lacerta schreiberi, Lacerta viridis (=L. bilineata) ein. Demnach ist das ontogenetische Wachstum von Kopf und Rumpf bei männlichen Tieren isometrisch, während Weibchen hingegen ein allometrisches Wachstum des Rumpfes aufweisen. Mit der verwendeten Methodik (Felsenstein) ließ sich der Geschlechtsdimorphismus am besten durch den weiblichen Fortpflanzungsaufwandes (u. a. Gelegegröße) und damit durch die überproportionale Rumpflänge weiblicher Tiere erklären. Bei Arten mit geringer Fruchtbarkeit bzw. konstanter Gelegegröße (hier: Podarcis hispanica) besteht hingegen nur ein geringer Selektionsdruck in Richtung auf große Weibchen. Dann können Männchen zum größten Geschlecht werden. Ebenfalls durchgeführte Größenmessungen an Beutetieren (Magen-Darminhalte) ergaben vergleichsweise gering abweichende Einnischungen männlicher und weiblicher Eidechsen. Eine Analyse der Mechanismen, die zur Ausprägung von Geschlechtsdimorphismen beitragen, ist sicherlich verdienstvoll. Allerdings ist fraglich, ob die hier verwendeten Methoden allein zu einer befriedigenden Klärung führen. Zum einen wurden pro Art nur wenige Individuen (zwischen 22 und 90) untersucht, was - auch nach Ansicht des Autors - statistisch unbefriedigend ist. Wichtiger dürfte aber wohl sein, dass keinerlei Hinweise darauf vorliegen, inwieweit die aus einem relativ begrenzten Gebiet stammenden Freilandfänge als “typisch“ für die Art gelten können. Hinzu kommt, dass sich mit der verwendeten Methode nur die Parameter weiblicher Fortpflanzungsaufwand und unterschiedliche Nahrungseinnischung, nicht jedoch der dritte vom Autor angesprochene Parameter “Kämpfe zwischen den Männchen“ untersuchen lässt. Zumindest in diesem Kontext bedarf es weiterer Daten, die nur durch Verhaltensbeobachtungen lebender Eidechsen zu gewinnen sind.
Braña, F. & Arrayago, M.J. & Bea, A. & Barahona, A. (1990) -
Male reproductive and fat body cycles were studies both in a mountain and a lowland population of the lizard Lacerta monticola cantabrica. Specimens were collected at two localities in Asturias (Northern Spain): Vegadeo (100 m a.s.l.) and Aramo (1450 m a.s.l.). General patterns of spermatogenesis in both populations were similar, and agree with the mixed type proposed by Saint Girons (1963). Spermatogenesis and testicular growth start in August and spermiogenesis begins in early autumn; before hibernation (October) spermiogenesis ceases earlier in the mountain than in the lowland population. The culminative phase of spermatogenesis and the testis weight peak occur after the onset of spring activity; there is then a progressive decrease that lasts until the end of the female`s ovulation cycle in July. Recrudescence of epididymides occurs mainly in late winter-early spring, with a pronounced development just after emergence from wintering; there is a delay in the epididymis peak of the mountain population. Both populations show an abrupt epididymal decline in the summer, between June and August. Fat bodies begin to increase in summer, just after the end of the reproductive period, reaching maximum weights in autumn prior to wintering. Pre-hibernation fat body weights for males at the two elevations are nearly the same (about 2% of body weight on average), but winter consumption is much larger in the lowland population in which there is some activity during this period. Fat bodies are not depleted at emergence and the remaining stores are utilized during the spring reproductive period, leading to minimum levels in June in both populations.
Braña, F. & González, F. & Barahona, A. (1992) -
Brändle, M. & Rödel, M.-O. (1994) -
Within ten years, data on twenty amphibian and thirty reptile species from the Iberian Peninsula were collected. The excursions were mainly centred on the northeast of Spain and the Extremadura. We report ob- servations on Salamandra salamandra almanzoris MÜLLER & HELLMICH, 1935, Rana pereti SEOANE, 1885, Lacerta viridis (LAURENTI, 1768), Lacerta monticola BOULENGER, 1905, and Malpolon monspes- sulanus (HERMANN, 1804). Additionally, all herpetological records and brief descriptions of the correspond- ing habitats are listed.
Breg, A. & Janota, B. & Peganc, M. & Petrovič, I. & Tome, S. & Vamberger, M. (2010) -
Brelih, S. (1954) -
Brelih, S. (1962) -
Brelih, S. & Džukić, G. (1974) -
Bressi, N. (1999) -
The Herpetological Collection of the Trieste Natural History Museum has almost 700 specimens of European Sauria belonging to about 33 taxa, including all the lacertids of northern Adriatic re- gions. The origin of the collection dates back to the foundation of the Museum of Trieste in 1846, but it was increased mainly between 1871 and 1939. The Herpetological Collection of the Trieste Natural History Museum has a great scientific and historical importance; it documents the varia- tion of the composition and distribution of northern Adriatic Sauria fauna during this century, in- cluding endemic varieties typical of little islands.
Brown, R.P. (2005) -
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.
Brown, R.P. & Perez-Mellado, V. (1993) -
The rock lizard Lacerta monticola is a climatic relict confined to three mountain ranges and one coastal region of the Iberian peninsula. Scalation was studied in specimens from localities encompassing all major parts of its range. Population differentiation was analysed using analyses of variance, z-transformations of within-locality character-correlations, canonical variate analyses, principal components analysis, and matrix association tests. Substantial differentiation was found between Pyrenees specimens and those from other localities, supporting a recent study which indicated that the Pyrenees populations could represent a different species. Considerable geographic variation is also found among populations from the remaining parts of its range, the pattern of which is not compatible with existing racial categories designated for this species.
Bruno, S. (1986) -
Bruno, S. & Maugeri, S. (1976) -
Busack, S.D. & Salvador, A. & Bauer, A.M. & Kaiser, H. (2016) -
A doctoral thesis presented in the late 1990s by óscar J. Arribas has been frequently cited as the authority for the nomenclatural status of two generic names in the lizard family Lacertidae, Darevskia and Iberolacerta. These names were again introduced, along with the addition of the subgeneric name Pyrenesaura, in 1999 via publication in an international, peer-reviewed journal. We discuss nomenclatural aspects of how these taxon names were presented, detail inconsistencies in the use of publication dates associated with these names, and resolve nomenclatural problems by demonstrating that the date of their first publication, in compliance with the International Code of Zoological Nomenclature, was 1999. This renders the name Darevskia Arribas, 1999 a junior invalid synonym of Caucasilacerta Harris, Arnold & Thomas, 1998, which, contrary to previous assessment, is not a nomen nudum, and must replace Darevskia as the valid name for this genus.
Cabela, A. & Grillitsch, H. & Happ, H. & Happ, F. & Koller, R. (1992) -
Cabela, A. & Grillitsch, H. & Tiedemann, F. (2002) -
Cabela, A. & Grillitsch, H. & Tiedemann, F. (2004) -
Cabela, A. & Grillitsch, H. & Tiedemann, F. (2007) -
Im Untersuchungsgebiet in den Lienzer Dolomiten (Kärnten, Österreich) kommen Iberolacerta horvathi (MÉHELY, 1904) und Podarcis muralis (LAURENTI, 1768) zwischen 714 und 1285 m ü. M. sympatrisch, zwischen 919 und 1057 m syntop vor. Rund 10 % der Eidechsenstandorte sind von beiden Arten etwa 50 % bzw. 40 % nur von P. muralis bzw. I. horvathi besetzt. Die beobachteten Individuendichten liegen bei I. horvathi mehrfach höher als bei P. muralis. Die felsig-steinigen Eidechsenhabitate – vom glatten, steilen Fels bis zur porösen, flachen Schutthalde – wurden nach ihrer Kompaktheit, Höhe, Neigung, Spaltenbildung und Oberflächenstruktur vier Typen zugeordnet, zu deren Besiedlung unterschiedliche Kletterfähigkeiten erforderlich sind. Auf dem glatten steilen Felsformationen kam nur I. horvathi vor, auf Schuttflächen nahezu ausschließlich P. muralis, Übergangsformen konnten beide Arten beherbergen.
Cabido, C. (2009) -
Cabido, C. & Galán, P. & López, P. & Martín, J. (2009) -
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.
Cabido, C. & Gonzalo, A. & López, P. & Martin, J. (2008) -
Cabido, C. & Iglesias-Carrasco, M. (2014) -
Cabido, C. & López, P. & Martin, J. (2006) -
Cabral, M.J. & Almeida, J. & Almeida, P.R. & Dellinger. T. & Almeida, N. de & Oliveira, M.E. & Palmeirim, J.M. & Queiroz, A.I. & Pogado, L. & Santos-Reis, M. (2005) -
Cafuta, V. (2010) -
Capizzi, D. (1999) -
The dietary habits of Lacerta horvathi MÉHELY, 1904, from the Tarvisio Forest (Carnic Alps, Province of
Udine) were studied using faecal pellet analysis. 81 specimens were analyzed for prey remnants in tlieir faecal pellets.
The sexes did not differ significantly in terms of SVL (one-way ANOVA: P < 0.30), but males were larger than females.
Lacerta horvathi fed mainly upon Opiliones and Araneae, while the proportion of insects was quite low. Both
mean number of prey items per faecal pellet and food niche breadth was higher in males than in females and juveniles,
but the differences between the three classes were not statistically significant. Compared with several other lacertid
species, L. horvathi seems to be much less a food generalist. In fact, it is strongly specialized in preying upon grounddwelling
arthropods. Furthermore, the prey types eaten suggest that this lizard is an active forager, as expected in
small-sized rupicolous lacertids.
Capizzi, D. & Luiselli, L. & Vignoli, L. (2006) -
Capizzi, D. & Luiselli, L. & Vignoli, L. (2007) -
Flight initiation distance in relation to substratum type, sex, reproductive status and tail condition was studied in two lacertid lizards with contrasting habits: the ground-dwelling common lizard Zootoca vivipara and the rupicolous Horvath`s rock lizard Iberolacerta horvathi. These species were studied in sympatric populations in a mountain area in North-Eastern Italy, Tarvisio Forest. Mean escape distance was significantly higher in I. horvathi than in Z. vivipara. In both species there were significant differences between sexes, with males escaping at longer distances than females but there were no significant differences between adults and subadults. In both species there were no differences in escape distance of females in different reproductive states. In Z. vivipara specimens with broken tails escaped at a shorter distance than individuals with intact tails. Substratum type had a significant effect on escape distance in both species.
Capula, M. & Lapini, L. & Capanna, E. (1989) -
The chromosomes of Lacerta horvfthi have been studied by means of conventional, C-banding, and silver-NOR techniques. The karyotype of this species, characterized by 36 acrocentric macrochromo- somes, lacks the typical pair of microchromosomes shared by all other lacertid lizards. It is hypothesized that the microchromosomes could have been translocated to the large elements of the karyotype. The occurrence of such a rearrangement in the chromosome complement ofL. horvhthi underlines its isolation from the other species of the subgenus Archaeolacerta. The C-banding analysis evidences the existence of a female sex heteromorphism in which the W-chromosome has the same shape and size of the Z, but differs from it in being completely heterochromatic. The nucleolar organizer regions (NORs) are located on a pair of medium size chromosomes in subtelomeric position, where the standard Giemsa-staining reveals secondary constrictions.
Capula, M. & Luiselli, L. (1990) -
Capula, M. & Luiselli, L. (1993) -
Capula, M. & Scalera, R. (1998) -
Carbonero, J. & Garcia-Diaz, P. & Ávila, C. & Arribas, O. & Lizana, M. (2016) -
seven species of the rock lizard genus Iberolacerta are represented on the iberian Peninsula. These lizards are considered to be under threat of extinction, with Iberolacerta martinezricai (ArribAs, 1996), being one of the most endangered reptiles in Europe. There is however, lack of knowledge about its conservation status, distribu- tion and ecology.
The authors’ surveys in 2007 and 2008 aimed at clarifying the distribution status of the species in Central spain, where 63 uTM squares (1 km x 1 km in size) were sampled and the density of the lizards, their distribution and habitat preferences studied. Iberolacerta martinezricai was detected in 23 out of the 63 uTM squares (36.5 %), with densities ranging from 25 to 50 lizards/ha. The statistical analysis revealed that the presence of the rock lizards was associated with altitude, lichen cover and rock/boulder size. Accordingly, the species is restricted to rocky slopes of the Peña de Francia mountain range. The results clearly indicate that the range of the species is highly restricted, that the habitats occupied are very specific, and that the population size is low compared with other species of the genus Iberolacerta. based on these findings, the species is categorized as Critically Endangered (Cr) under the criteria of the international union for the Conservation of nature (iuCn).
Carranza, S. & Arnold, E.N. & Amat, F. (2004) -
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.
Carrascal, L.M. & López, P. & Martin, J. & Salvador, A. (1992) -
This paper presents an observational and experimental study of the basking behaviour and heat exchange rate of the montane lizard Lscerta monticola. The results obtained by these procedures were coupled in order to understand behavioural mechanims promoting effective thermoregulation at high altitudes. Heating rate was higher when body size was smaller, and substrate temperature and sun rays incidence angle were higher. The lizards cooled faster when body size and substrate temperature were lower, and when the body temperature of the lizard going into shadow was higher. Time exposed to sun and mean duration of basking periods were longer early in the morning, while bask frequency increased through the morning. Our results suggest that time devoted to basking is mainly obtained by regulating bask duration. Lizards obtained the necessary time for heating by means of long basking periods. Mean travel distance per minute and distance to the nearest refuge increased from early morning to midday. These behavioural variables were tightly correlated with the expected heating rate of individuals. Body size affects thermoregulatory behaviour as well as locomotor activity. Juvenile lizards, with small body mass and high surface-to-volume ratios, were subjected to faster heating and cooling rates, basked more frequently than adults (but during shorter periods), and devoted more time to locomotion than adults. The thermoregulatory behaviour of L. monticola is the result of the combination of shuttling heliothermy by basking and the exploitation of thermal opportunities offered by patches in shade through thermal exchange with the substrate.
Carretero, M.A. (2004) -
Lacertids are the dominant group of lizards throughout the Mediterranean Basin. Their role in food web transfer of matter and energy from arthropods and other small invertebrates to birds and mammals constitutes a major function within Mediterranean ecosystems. For many years, prey consumption by lacertids was thought to be almost indiscriminate, not much more than a byproduct of habitat use. However, increasing evidence does not support this passive view. Analyses of prey availability have revealed active prey selection/avoidance in several species. Others show an internal tendency (i.e., historical constraints) to consume specific animal items (ants, clumped prey) or plant matter (seeds, nectar, pollen, leaves). Behavioural experiments showed that lacertids not only identify different prey types by both visual and chemical cues but also modify their feeding behaviour integrating past experiences. Furthermore, size, sex, reproductive state, body condition, tail loss and probably other lizard features are relevant for feeding ecology. However, less attention has been devoted to abiotic factors such as temperature and humidity. More experimental studies of the influences of competitors, predators and parasites on diet are needed. Even though it is controversial, optimal foraging theory provides a conceptual background for future studies. The evolutionary history of the various lacertid lineages, which constrains their morphology and physiology and eventually produces exaptative traits, is to be considered as well. Finally, methodology in field sampling, lab work and statistical analysis needs to be developed. Recommendations are given as to when and where to sample, which compartment should be analysed, which is the appropriate sample size, how to assess trophic availability, which statistical descriptors should be used and how they should be compared.
Carretero, M.A. (2018) -
Carretero, M.A. & Martínez-Solano, Í. & Ayllón, E. & Llorente, G. (2018) -
Cassol, M. & Romanazzi, E. & Cerbo, A.R. di & Vettorazzo, E. (2016) -
Cassol, M. & Romanazzi, E. & Cerbo, A.R. di & Vettorazzo, E. (2017) -
Castilla, A.M. & Van Damme, R. & Bauwens, D. (1999) -
We discuss three aspects of the thermal biology of lacertid lizards. First, we provide an overview
of the available data on field body temperatures (Tb), the thermal sensitivity of various performance
functions and selected body temperatures in different species of lacertid lizards. We also
briefly summarise information on the mechanisms of thermoregulation. Second, we discuss recent
developments to estimate the »precision« of thermoregulation, and the contribution of distinct behavioural
mechanisms. Finally, we revise available evidence for the existence of evolutionary adjustments
of thermal characteristics in lacertid lizards. Existing studies have mainly dealt with
within- and among-species differences in thermoregulatory behaviour (selected temperatures) and
thermal physiology of adults (optimal temperatures, heating rates). Available data provide only
limited evidence for clear-cut evolutionary shifts in thermal physiology characteristics along climatic
Ceia-Hasse, A. & Sinervo, B. & Vicente, L.& Pereira, H.M. (2014) -
Uncertainty in projections of global change impacts on biodiversity over the 21st century is high. Improved predictive accuracy is needed, highlighting the importance of using different types of models when predicting species range shifts. However, this is still rarely done. Our approach integrates the outputs of a spatially-explicit physiologically inspired model of extinction and correlative species distribution models to assess climate-change induced range shifts of three European reptile species (Lacerta lepida, Iberolacerta monticola, and Hemidactylus turcicus) in the coming decades. We integrated the two types of models by mapping and quantifying agreement and disagreement between their projections. We analyzed the relationships between climate change and projected range shifts. Agreement between model projections varied greatly between species and depended on whether or not they consider dispersal ability. Under our approach, the reliability of predictions is greatest where the predictions of these different types of models converge, and in this way uncertainty is reduced; sites where this convergence occurs are characterized by both current high temperatures and significant future temperature increase, suggesting they may become hotspots of local extinctions. Moreover, this approach can be readily implemented with other types of models.
Clemente, J.J.P. (2005) -
Cocca, W. & Žagar, A. & Carretero, M.A. & Krofel, M. & Luznik, M. & Podnar, M. & Tvrtkovic, N. & Crottini, A. (2016) -
The Horvath’s rock lizard, Iberolacerta horvathi, is occurring from 200 to 2000 m a.s.l. on the eastern Alps (NE Italy, S Austria, NW Slovenia) and the northern Dinaric Mountains (central and S Slovenia and NW Croatia). This species is the only representative of Iberolacerta outside Iberian Peninsula and currently shows a discontinuous occupational range. We collected tissue samples and extracted total genomic DNA of 110 samples from 23 localities across its entire distributional range. We grouped localities into three main geographical areas: northwestern (NW), central and southeastern (SE). For all samples we amplified a fragment of the mtDNA cytochrome b (Cytb) gene whereas a fragment of the nuclear melanocortin 1 receptor (MC1R) gene was amplified for a subset of 53 samples. Despite the relatively narrow geographical scale, the Cytb haplotype network indicated a high number of unique haplotypes present in each area (6 in the NW and 14 in the central and SE). In NW and in central area we detected two shared haplotypes: a) the widely distributed H5, and b) the H21, shared by only two populations. On the contrary, SE populations did not share any Cytb haplotype with other areas. The nuclear haplotype network analysis gave similar results, where multiple private alleles occurred in each area (4 in NW and central, and 3 in SE) and a widespread haplotype shared across the three areas. Results suggest that current distributional fragmentation is partially mirrored by the phylogeographic history of the species, with private alleles occurring in each of the main geographic areas, indicating limited gene flow among populations at least in the recent past. Knowledge of genetic structure of this endemic species provides essential information to orient activities to ensure its future conservation.
Cooper Jr., W.E. & López, P. & Martín, J. & Pérez-Mellado, V. (2012) -
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.
Cooper Jr., W.E. & Martin, J. & Lopez, P. (2003) -
Cooper Jr., W.E. & Pyron, A. & Garland, T. jr. (2014) -
One of Darwin`s most widely known conjectures is that prey are tame on remote islands, where mammalian predators are absent. Many species appear to permit close approach on such islands, but no comparative studies have demonstrated reduced wariness quantified as flight initiation distance (FID; i.e. predator–prey distance when the prey begins to flee) in comparison with mainland relatives. We used the phylogenetic comparative method to assess influence of distance from the mainland and island area on FID of 66 lizard species. Because body size and predator approach speed affect predation risk, we included these as independent variables. Multiple regression showed that FID decreases as distance from mainland increases and is shorter in island than mainland populations. Although FID increased as area increased in some models, collinearity made it difficult to separate effects of area from distance and island occupancy. FID increases as SVL increases and approach speed increases; these effects are statistically independent of effects of distance to mainland and island occupancy. Ordinary least-squares models fit the data better than phylogenetic regressions, indicating little or no phylogenetic signal in residual FID after accounting for the independent variables. Our results demonstrate that island tameness is a real phenomenon in lizards.
Corti, C. & Lo Cascio, P. (2002) -
Cox, N. & Chanson, J. & Stuart, S. (2006) -
Crespo, E.G. & Cei, J.M. (1975) -
Crochet, P.-A. & Chaline, O. & Surget-Groba, Y. & Debain, C. & Cheylana, M. (2004) -
Crochet, P.A. & Rufray, V. & Viglione, J. & Geniez, P. (1996) -
Cyrén, O. (1928) -
Cyrén, O. (1941) -
Dall`Asta, A. & Richard, J. (2016) -
Davies, R.S. (1984) -
Delibes, A. & Salvador, A. (1986) -
Delic, T. & Zagar, A. (2011) -
Deschandol, F. (2011) -
Dieckmann, M. (2018) -
Dieckmann, M. (2019) -
Diesener, G. & Reichholf, J. (1986) -
Dolce, S. & Lapini, L. (1987) -
Domènech, S. (1997) -
Dominguez, L. & Elvira, B. & Vigal, C.R. (1981) -
Dominguez, L. & Elvira, B. & Vigal, C.R. (1982) -
Džukić, G. & Kalezić, M. (2004) -
Dzukic, G. & Kalezic, M.L. (2001) -
Elvira, B. & Palacios, F. (1981) -
Elvira, B. & Vigal, C.R. (1982) -
Elvira, B. & Vigal, C.R. (1985) -
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.
Ernst, K. (2017) -
Das Ziel dieser Studie ist, neue Erkenntnisse über die bevorzugten ökologische Nischen und die aktuelle Verbreitung der Kroatischen Gebirgseidechse Iberolacerta horvathi (MÉHELY, 1904) innerhalb Österreichs zu gewinnen. Eine aussagekräftige Vorhersage werden wir mittels Modellierung der potentiellen Verbreitung (SDM – Species‘ distribution modeling) für Österreich treffen und darstellen können. Für die korrelative Modellierung implementieren und analysieren wir artspezifische Habitat-Präferenzen unter Berücksichtigung geologischer, topografischer, bioklimatischer und ökologischer Variablen, basierend auf Daten bisher bekannter Fundorte. Eine anschließende Freilandstudie an die vom Modell berechneten Standorte mit hoher Wahrscheinlichkeit weiterer Vorkommen, soll schlussendlich zur Entdeckung noch unbekannter besetzter Nischen von I. horvathi in Österreich führen. Endresultat der Studie wird eine umfassende Art-Charakterisierung und eine Verbreitungskarte der potentiellen, als auch realisierten Habitate, entlang der nördlichen Verbreitungsgrenze der Kroatischen Gebirgseidechse in Österreich sein. Außerdem testen wir die Aussagekraft der Modellierung, die zukünftig auch als hilfreiche, flexible Methode in Bezug auf Artenschutz Anwendung finden könnte.
Ernst, K. & Gollmann, G. & Plutzar, C. & Schweiger, S. (2017) -
Faberl, F. & Faberl, H. (1991) -
Ferchaud, A.L. & Eudeline, R. & Arnal, V. & Cheylan, M. & Pottier, G. & Leblois, R. & Crochet, P.-A. (2015) -
Historical factors, current population size, population connectivity and selective processes at linked loci contribute to shaping contemporary patterns of neutral genetic diversity. It is now widely acknowledged that nuclear and mitochondrial markers react differently to current demography as well as to past history, so the use of both types of markers is often advocated to gain insight on both historical and contemporary processes. We used 12 microsatellite loci genotyped in 13 populations of a mountain lizard (Iberolacerta bonnali) to test whether the historical scenario favoured by a previous mitochondrial study was also supported by nuclear markers and thereby evaluated the consequences of postglacial range movements on nuclear diversity. Congruent signals of recent history were revealed by nuclear and mitochondrial markers using an Approximate Bayesian computation approach, but contemporary patterns of mtDNA and nuclear DNA diversity were radically different. Although dispersal in this species is probably highly restricted at all spatial scales, colonization abilities have been historically good, suggesting capability for reestablishment of locally extinct populations except in fully disconnected habitats.
Fernández, J. & Arribas, O. (2014) -
A male of Aurelio’s Rock Lizard was observed devouring its own distal part of the tail after suffering an incomplete amputation (autourophagy). This is the first observed case of this phenomenon and is compared with other cannibalism observations in Sauria.
Franzen, M. & Gruber, H.-J. (2004) -
Franzen, M. & Gruber. H.-J. & Heckes, U. (1993) -
Fretey, J, (1986) -
Fugas, M.C. & Caeiro, F. (2012) -
Amoeba associated microorganisms (ARMs) are bacteria or viruses that share a symbiotic relationship with amoebas. Many ARMs are associated with human diseases and it has been reported the acquisition of resistance inside their host. These facts highlight the importance of finding and characterizing ARMs in a public health’s perspective. In the present work, amoebas from environmental samples have been isolated and identified by optical microscopy and molecular methods. Amoeba associated viruses, especially from the NCLDV group, were also subjected to a PCR screening. From the 12 collected samples, it was possible to isolate 17 amoebas, of which 11 were identified by molecular methods. No viruses were detected but in 2 of these amoebas, a bacterium was identified which shares an 89% sequence identity with the Marivirga tractuosa species. If amoebas allow the replication of a Mimivirus, which belongs to the NCLDV group, would they be able to replicate iridoviruses that belong to the same group? An Acanthamoeba castellanii cell line was experimentally inoculated with three Ranavirus: FV3 (Frog Virus 3), Ma3B and BoA (respectively, a Triturus marmoratus and a Triturus boscai isolate). The PCR test suggested an association between A. castellanii cells with the Ma3B and BoA viruses. VLPs (Virus-like particles) were visualized by electron microscopy in the Ma3B inoculated amoebas. Ranavirus from several Triturus species are currently being characterized and the sequences of their Major Capsid Protein and DNA polymerase genes are identical. Could the sequencing of a less conserved gene like the Flap endonuclease, allow the distinction? The sequencing and analysis of the Flap gene was able to group viruses FV3 and LMO (Lacerta monticula virus) as they present an equal gene sequence. The three Triturus marmoratus isolates presented an equal Flap gene amongst them and were also grouped. Between the two groups, this gene sequence has a difference of 8 nucleotides, confirming the separation of the FV3 and LMO viruses from the other iridoviruses tested.
Gabirot, M. & Balleri, A. & López, P. & Martin, J. (2000) -
Research on fluctuating asymmetry (FA)-mediated sexual selection has focused almost exclusively on visual signals and ignored chemical communication despite the fact that many species rely on chemical signals for attracting mates. Female mate choice based on visual traits appears to be rare in lizards. However, the femoral glands of male lizards produce pheromones which might transmit chemical information about an individual`s developmental stability. Therefore, we hypothesized that mate choice may be based on chemical cues. We analysed the effect of the developmental stability levels of males on the attractiveness of males` scents to females in a laboratory experiment with the lizard Lacerta monticola. When we offered two males of similar body size, females preferentially associated with the scents of males with low FA in their femoral pores and also with the scents of males with a higher number of femoral pores. This suggested that the females were able to discriminate the FA of the males by chemical signals alone and that the females preferred to be in areas marked by males of high quality, thus increasing their opportunities of mating with males of high quality. We suggest that the quality and/or amount of male pheromones could communicate the heritable genetic quality of a male to the female and thereby serve as the basis for adaptive female choice in lizards.
Galán Regalado, P. (1982) -
Galán Regalado, P. (1991) -
Galan, P. (1982) -
Galan, P. (1983) -
Galan, P. (1992) -
Galán, P. (1999) -
Galan, P. (2005) -
The main threats to the amphibians and reptiles of Galicia (NW Spain) are studied in relation to the information obtained during the last 30 years (1975-2005). The most negative factors affecting amphibians and reptiles in this territory are the human alteration of both aquatic and terrestrial habitats, and the introduction of exotic species (mainly freshwater crustaceans and fishes). These menaces have led to the extinction of some Galician populations of amphibian and reptiles. Other factors, such as emergent diseases, can be very important, but they have not been studied yet. This last threat is one of the main causes for the global amphibian decline. The recent status of the Galician herpetofauna is also analysed. According to the main population declines and areal reduction observed, the most endangered amphibian species is Pelobates cultripes. Other amphibians, such as Chioglossa lusitanica, Rana iberica and Rana temporaria, are also importantly put at risk. The most endangered reptile species is Emys orbicularis. The restricted and limited Galician populations of Lacerta vivipara, Chalcides bedriagai and Vipera latasti are also threatened. The available information for other reptile species (Blanus cinereus, Psammodromus hispanicus, etc.), is still very scarce, but it is possible that their situation is also precarious. The particular status of the insular populations of amphibians and reptiles from the islands off the Galician coast is also analysed.
Galán, P. (2006) -
Galán, P. (2008) -
Changes in the coloration of the lacertid lizards Iberolacerta monticola and Podarcis bocagei with age in populations from NW Spain are described. The onset of sexual maturity in P. bocagei males involves a change in the ventral (yellow) and dorsal (green) colorations, which is diﬀerent from immature males (dorsally brownish in color). In I. monticola males, the ventral coloration also changes to a deep green when they reach maturity, while the dorsal coloration remains brownish as in the immature specimens. In this species, the green dorsal coloration is acquired gradually after maturity. Only the oldest individuals have a predominantly green dorsal coloration. The diﬀerences between the two species in the time males take to acquire the green dorsal coloration could be related to their diﬀ erent longevity. The coloring is acquired gradually in the most long-lived species ( I. monticola ). A ﬁeld study was carried out on the behaviour of adult males of I. monticola during the reproductive period. The males with green dorsal coloration were seen to pair with females signiﬁ cantly more frequently than those with the brownish dorsal color. The increase in the green dorsal coloration (conspicuous) with the size and age of the males of this species would appear to have a clear function as an intersexual or intrasexual signal.
Galan, P. (2010) -
Galan, P. (2011) -
Galán, P. (2011) -
Galan, P. (2012) -
Galan, P. (2019) -
Galán, P. & Arribas, O. (2007) -
Galán, P. & Ferreiro, R. & Naviera, H. F. (2007) -
Galán, P. & Remón, N. & Vila, M. & Fernández, E. & Naveira, H. (2006) -
Galán, P. & Vila, M. & Remón, N. & Naveira, H.F. (2007) -
The Iberian rock lizard (Iberolacerta monticola) has gone extinct or severely declined in several low-altitude and isolated populations in Galicia during the last three decades. In order to assess the conservation status of this species in NW Iberia, we recorded morpho- and eco- logical data of individuals from eleven localities of this area. In addition, the genetic variability of these specimens was screened by combi- ning nuclear (microsatellite) and mitochondrial markers (470 bp from the control region). The population established at O Pindo (A Coruña), at least 100 km apart from other populations, shows a conspicuous diagnostic difference in its mitochondrial lineage and the largest diffe- rentiation at microsatellite loci, as well as some morphological differentiation most likely linked to the particular habitat of this site, a relatively arid and rocky environment.
Gállego-Castejón, L. & López, S. (1983) -
Garcia-Diaz, P. (2011) -
Garcia-Diez, T. & González-Fernández, J.E. (2013) -
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.
Garcia-Porta, J. & Irisarri, I. & Kirchner, M. & Rodríguez, A. & Kirchhof, S. & Brown, J.L. & MacLeod, A. & Turner, A.P. & Ahmadzadeh, F. & Albaladejo, G. & Crnobrnja-Isailovic, J. & Riva, I. de la & Fawzi, A. & Galán, P. & Göçmen, B. & Harris, D.J. & Jiménez-Robles, O. & Joger, U. & Jovanović Glavaš, O. & Karış, M. & Koziel, G. & Künzel, S. & Lyra, M. & Miles, D. & Nogales, M. & Oğuz, M.A. & Paf (2019) -
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.
Garcia-Roa, R. (2017) -
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.
García-Roa, R. & Iglesias-Carrasco, M. & Garin-Barrio, I. & Cabido, C. (2015) -
Garcia-Roa, R. & Ortega, J. & López, P. & Martin, J. (2015) -
Garcia-Roa, R. & Sáiz, J. & Gómara, B. & López, P. & Martin, J. (2017) -
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.
Garcia-Roa, R. & Sáiz, J. & Gómara, B. & López, P. & Martin, J. (2018) -
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.
Garcia-Roja, R. & Llusia, D. & López, P. & Martin, J. (2017) -
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.
Giovannotti, M. & Rojo, V., & Nisi-Cerioni, P. & González-Tizón, A. & Martinez-Lage, A. & Splendiana, A. & Naveira, H. & Ruggeri, P. & Arribas, O. & Olmo, E. & Caputo Barucchi, V. (2013) -
Satellite DNAs represent a large portion of all high eukaryotic genomes. They consist of numerous very similar repeated sequences, tandemly arranged in large clusters up to 100 million base pairs in length, usually located in the heterochromatic parts of chromosomes. The biological significance of satDNAs is still under discussion, but most of their proposed functions are related to heterochromatin and/or centromere formation and function. Because information about the structure of reptilian satDNA is far from exhaustive, we present a molecular and cytogenetic characterization of two satDNA families in four lacertid species. Two families of tandemly repeated DNAs, namely TaqI and HindIII satDNAs, have been cloned and sequenced from four species belonging to the genus Iberolacerta. These satDNAs are characterized by a monomer length of 171–188 and 170–172 bp, and by an AT content of 60.5% and 58.1%, respectively. FISH experiments with TaqI satDNA probe produced bright signals in pericentromeric regions of a subset of chromosomes whereas all the centromeres were marked by HindIII probe. The results obtained in this study suggest that chromosome location and abundance of satDNAs influence the evolution of these elements, with centromeric families evolving tenfold faster than interstitial/pericentromeric ones. Such different rates render different satellites useful for phylogenetic investigation at different taxonomic ranks.
Gisbert, J. (1981) -
Glandt, D. (2010) -
Glandt, D. (2011) -
Godinho, R. & Teixeira, J. & Rebelo, R. & Segurado, P. & Loureiro, A. & Alvares, F. & Gomes, N. & Gardoso, P. & Camilo-Alves, C. & Brito, J.C. (1999) -
Records for the amphibian and reptile species of continental Portugal were plotted on maps using the 10 10 km squares of the UTM grid system. A total of 9394 observations were collected, from which 6485 re- cords came from previous national atlases, 1790 from recent published and unpublished work, and 1119 co- rrespond to new observations made by the authors and their collaborators. Overall 31.0% of the records are pre- sented for the first time in a national atlas. Remarkable new records include those for Tarentola mauritanica and Blanus cinereus, enlarging their known range into areas where they were not thought to occur. We present and discuss amphibian and reptile species-density maps.
Grillitsch, H. & Cabela, A. (1992) -
The attempt is made to define the distributional patterns of the reptiles in Carinthia (Austria) by selected parameters of climat and vegetation.
Grillitsch, H. & Tiedemann, F. (1986) -
Günther, R. (1996) -
Harris, D.J. & Arnold, E.N. & Thomas, R.H. (1998) -
DNA sequences from parts of the 12S, 16S and cytochrome b mitochondrial genes, which totalled 1049 aligned base pairs, were used to estimate the relationships of 49 species of Lacertidae, including representatives of 19 out of the 23 recognized genera and 23 species of the paraphyletic genus Lacerta. These data were used, together with morphological information, to estimate the relationships within the family. Molecular evidence corroborates the monophyletic status of many genera and species groups originally based on morphology. It indicates that Psammodromus forms a clade with Gallotia, which is the sister taxon of all other lacertids. These comprise three units: the primarily Afrotropical armatured group; the largely Oriental Takydromus; and the west Palaearctic Lacerta and its derivatives, Podarcis and Algyroides. Morphology also supports the first three assemblages, but suggests that they are derived from a paraphyletic Lacerta. Within Lacerta and its allies, DNA sequence analysis corroborates the affinity of some members of each of the subgenera Lacerta s. str. and Timon, and of the L. saxicola group. It also supports the relationship of L. monticola, L. bonnali and L. horvathi, and suggests that the L. parva–L. fraasi clade and L. brandti are not related to Psammodromus–Gallotia, as morphology indicates, but instead are associated respectively with the L. danfordi and L. saxicola groups. DNA sequence data provide additional evidence that the eastern Arabian `Lacerta` jayakari and `L.` cyanura are members of the armatured clade and also sister species. Our analysis supports an origin for present lacertids in west Eurasia. The armatured clade invaded Africa, probably in the mid-Miocene, spreading widely and evolving increasingly xeric-adapted forms, one lineage of which later moved back into the Palaearctic. `Lacerta` jayakari and `L.` cyanura are assigned to Omanosaura, Lutz and Mayer 1986. The name Gallotiinae Cano, Baez, Lopez-Jurado and Ortega, 1984 is available for the Gallotia–Psammodromus clade, Eremiainae Shcherbak 1975 for the armatured clade and Lacertinae for Lacerta, Podarcis and Algyroides. Two new subgenera of Lacerta are proposed here: Caucasilacerta for L. saxicola and its allies, and Parvilacerta for L. parva and L. fraasi.
Hassl, A.R. (2012) -
Helmich, W. (1935) -
Henle, K. & Dick, D. & Harpke, A. & Kühn, I. & Schweiger, O. & Settele, J. (2008) -
Henle, K. & Steinicke, H. & Gruttke, H. (2004) -
Herrando-Pérez, S. & Ferri-Yáñez, F. & Monasterio, C. & Beukema, W. & Gomes, V. & Belliure, J. & Chown, S.L. & Vieites, D.R. & Araújo, M.B. (2019) -
1. Research addressing the effects of global warming on the distribution and persistence of species generally assumes that population variation in thermal tolerance is spatially constant or overridden by interspecific variation. Typically, this rationale is implicit in sourcing one critical thermal maximum (CTmax) population estimate per species to model spatiotemporal cross‐taxa variation in heat tolerance. Theory suggests that such an approach could result in biased or imprecise estimates and forecasts of impact from climate warming, but limited empirical evidence in support of those expectations exists. 2. We experimentally quantify the magnitude of intraspecific variation in CTmax among lizard populations, and the extent to which incorporating such variability can alter estimates of climate impact through a biophysical model. To do so, we measured CTmax from 59 populations of 15 Iberian lizard species (304 individuals). 3. The overall median CTmax across all individuals from all species was 42.8 °C and ranged from 40.5 to 48.3 °C, with species medians decreasing through xeric, climate‐generalist and mesic taxa. We found strong statistical support for intraspecific differentiation in CTmax by up to a median of 3 °C among populations. We show that annual restricted activity (operative temperature > CTmax) over the Iberian distribution of our study species differs by a median of > 80 hours per 25‐km2 grid cell based on different population‐level CTmax estimates. This discrepancy leads to predictions of spatial variation in annual restricted activity to change by more than 20 days for six of the study species. 4. Considering that during restriction periods, reptiles should be unable to feed and reproduce, current projections of climate‐change impacts on the fitness of ectotherm fauna could be under‐ or over‐estimated depending on which population is chosen to represent the physiological spectra of the species in question. Mapping heat tolerance over the full geographical ranges of single species is thus critical to address cross‐taxa patterns and drivers of heat tolerance in a biologically comprehensive way.
Hipsley, C. (2012) -
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.
Hipsley, C. & Himmelmann L. & Metzler D. & Müller J. (2009) -
Background: Although current molecular clock methods offer greater flexibility in modelling
evolutionary events, calibration of the clock with dates from the fossil record is still problematic
for many groups. Here we implement several new approaches in molecular dating to estimate the
evolutionary ages of Lacertidae, an Old World family of lizards with a poor fossil record and
uncertain phylogeny. Four different models of rate variation are tested in a new program for
Bayesian phylogenetic analysis called TreeTime, based on a combination of mitochondrial and
nuclear gene sequences. We incorporate paleontological uncertainty into divergence estimates by
expressing multiple calibration dates as a range of probabilistic distributions. We also test the
reliability of our proposed calibrations by exploring effects of individual priors on posterior
Results: According to the most reliable model, as indicated by Bayes factor comparison, modern
lacertids arose shortly after the K/T transition and entered Africa about 45 million years ago, with
the majority of their African radiation occurring in the Eocene and Oligocene. Our findings indicate
much earlier origins for these clades than previously reported, and we discuss our results in light
of paleogeographic trends during the Cenozoic.
Conclusion: This study represents the first attempt to estimate evolutionary ages of a specific
group of reptiles exhibiting uncertain phylogenetic relationships, molecular rate variation and a
poor fossil record. Our results emphasize the sensitivity of molecular divergence dates to fossil
calibrations, and support the use of combined molecular data sets and multiple, well-spaced dates
from the fossil record as minimum node constraints. The bioinformatics program used here,
TreeTime, is publicly available, and we recommend its use for molecular dating of taxa faced with
Hipsley, C.A. & Miles, D.B. & Müller, J. (2014) -
While global variation in taxonomic diversity is strongly linked to latitude, the extent to which morphological disparity follows geographical gradients is less well known. We estimated patterns of lineage diversification, morphological disparity and rates of phenotypic evolution in the Old World lizard family Lacertidae, which displays a nearly inverse latitudinal diversity gradient with decreasing species richness towards the tropics. We found that lacertids exhibit relatively constant rates of lineage accumulation over time, although the majority of morphological variation appears to have originated during recent divergence events, resulting in increased partitioning of disparity within subclades. Among subclades, tropical arboreal taxa exhibited the fastest rates of shape change while temperate European taxa were the slowest, resulting in an inverse relationship between latitudinal diversity and rates of phenotypic evolution. This pattern demonstrates a compelling counterexample to the ecological opportunity theory of diversification, suggesting an uncoupling of the processes generating species diversity and morphological differentiation across spatial scales.
Horváth, G. & Jiménez-Robles, O. & Martin, J. & López, P. & Riva, I. de la & Herczeg, G. (2020) -
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.
Horvath, G. & López, P. & Martin, J. (2014) -
Horváth, G. & Martin, J. & López, P. (2016) -
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.
Horváth, G. & Martin, J. & López, P. & Garamszegi, L.Z. & Bertok, P. & Herczeg, G. (2016) -
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.
Horváth, G. & Martín, J. & López, P. & Garamszegi, L.Z. & Herczeg, G. (2017) -
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.
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.
Huyghe, K. & Herrel, A. & Husak, J.F. & Damme, R. van (2007) -
Iglesias-Carrasco, M. & Cabido, C. (2016) -
Iglesias-Carrasco, M. & Head, M.L. & Martin, J. & Cabido, C. (2018) -
Environmental conditions experienced by a species during its evolutionary history may shape the signals it uses for communication. Consequently, rapid environmental changes may lead to less effective signals, which interfere with communication between individuals, altering life history traits such as predator detection and mate searching. Increased temperature can reduce the efficacy of scent marks released by male lizards, but the extent to which this negative effect is related to specific biological traits and evolutionary histories across species and populations have not been explored. We experimentally tested how increased temperature affects the efficacy of chemical signals of high- and low- altitude populations of three lizard species that differ in their ecological requirements and altitudinal distributions. We tested the behavioral chemosensory responses of males from each species and population to male scent marks that had been incubated at one of two temperatures (cold 16°C or hot 20°C). In high- altitude populations of a mountain species (Iberolacerta monticola), the efficacy of chemical signals (i.e., latency time and number of tongue flicks) was lower after scent marks had been exposed to a hot temperature. The temperature that scent marks were incubated at did not affect the efficacy of chemical signals in a ubiquitous species (Podarcis muralis) or another mountain species (I. bonalli). Our results suggest that specific ecological traits arising through local adaptation to restricted distributions may be important in determining species vulnerability to climatic change
Ineich, I. & Doronin, I. (2017) -
Ineich, I. & Doronin, I. & Lescure, J. (2017) -
The life and work of the Alsatian Louis Amédée Lantz (1886-1953), a pioneer of European herpetology. A trained and professional research-chemist, the Alsatian Louis Amédée Lantz was an active and particularly dynamic herpetologist. His residence of nearly twelve years in Russia enabled him to make several field trips to regions such as the Caucasus and Iran. He always carried with him a home made telescopic net that he used to catch lizards. That net became symbolic of Lantz for most people around him. He realized significant herpetological collections mainly deposited in the Museum of the Zoology Institute of the Russian Academy of Sciences in St. Petersburg. Soon after the Russian Revolution in 1917, he had to leave the country and stayed for a few years in France before joining England. During this period, Lantz visited the French Pyrenees and the Mediterranean border (Hyeres Islands, Bandol), enabling him to discover for the first time oviparous populations of common lizard and also to describe a new species of lizard from the French Pyrenees. Lantz is also at the origin of the discovery of unisexual reproduction by parthenogenesis in reptiles. His collaboration with the Swedish amateur herpetologist Otto Cyrén, whom he met through the German herpetologist Willy Wolterstorff, was particularly successful throughout his career. Lantz died at the age of 66 years, in Switzerland, as he was about to join the team of herpetologists at the Natural History Museum in Paris.
Ineich, I. & Doronin, I.V. & Cheylan, M. & Campbell, P.D. (2019) -
Several recent papers have reviewed the life and work of French herpetologist Louis Amédée Lantz. They have detailed the composition of his collections deposited in several museums. However, since then, several other important specimens from his collections deposited at the Natural History Museum (NHM, UK) have come to light and we here present all of them in detail. We discovered paralectotypes of Lacerta saxicola obscura Lantz & Cyrén (BMNH 19184.108.40.206–7), syntypes of Lacerta boemica Sukhov (BMNH 19220.127.116.11–30, BMNH 1965.337–342) and Lacerta viridis media Lantz & Cyrén (BMNH 1918.104.22.168, 1966.512). We also identified numerous specimens from the French Mediterranean islands in Lantz’s collection deposited at the NHM, some of which represent the first reported specimens of their species from certain islands. We here provide data on all these specimens. We also place the Mediterranean island specimens from the Lantz collection in their historical context and emphasize the historical and taxonomic value of these collections.
Janev-Hutinec, B. & Kletečki, E. & Lazar, B. & Podnar-Lešić, M. & Skejić, J. & Tadić, Z. & Tvrtković, N. (2006) -
Jelić, D. (2014) -
An updated checklist is presented for the amphibian and reptile fauna of Croatia. According to historical and recent literature sources, 20 amphibian and 41 reptile species have been recorded so far. Two reptile species, Blanus strauchi and Typhlops vermicularis, are considered doubtful, although there are individual museum vouchers for the country. Presence of a total of 59 species can be considered valid, with two additional considered doubtful. This paper summarizes data from 700 scientific and popular publications, which is the first attempt to publish a complete Croatian herpetological bibliography. Full bibliography is also given for all occurring species and constitutes a basis for future publications.
V članku je podan posodobljen seznam dvoživk in plazilcev Hrvaške. Po zgodovinskih in recentnih literaturnih virih je bilo doslej v državi zabeleženih 20 vrst dvoživk in 41 vrst plazilcev. Za dve vrsti plazilcev, Blanus strauchi in Typhlops vermicularis, je pojavljanje v državi vprašljivo, čeprav obstajajo njuni muzejski primerki. Tako je bilo skupaj najdenih 59 vrst, medtem ko sta dve vprašljivi. V članku je povzetih 700 strokovnih in poljudnih publikacij, kar je prvi poskus objave celotne bibliografije o herpetološki favni Hrvaške. Podana je tudi celotna bibliografija za vse vrste, ki je hkrati izhodišče za prihodnje objave.
Jelić, D. & Karaica, B. & Koren, T. & Sterijovski, B. & Dragičević,P. & Podnar Lešić, M. & Kovač Konrad, P. & Treer, D. & Kuljerić, M. & Peranić, I. (2015) -
Jelić, D. & Kuljerić, M. & Koren, T. & Treer, D. & Šalamon, D. & Lončar, M. & Podnar Lešić, M. & Janev Hutinec, B. & Bogdanović, T. & Mekinić, S. & Jelić, K. (2012) -
Jelic, D. & Kuljeric, M. & Koren, T. & Treer, D. & Salomon, D. & Loncar, M. & Podnar Lesic, M. & Hutinec, B.J. & Bogdanovic, T. & Mekinic, S. & Jelic, K. (2012) -
Jiménez-Robles, O. & de la Riva, I. (2019) -
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.
Kapli, P. & Poulakakis, N. & Lymberakis, P. & Mylonas, M. (2011) -
the Lacertidae is one of the most diverse and widespread lizard families throughout Eurasia and Africa. Several studies so far have attempted to unravel the phylogeny of Lacertidae using morphological and mole- cular data. However, the intra-family relationships remain unclear. in an effort to explore the phylogenetic relationships within the family Lacertidae, a concatenated dataset of 5727 bp from six genes (two nuclear and four mitochondrial) and 40 genera was assembled based on GenBank database. Phylogenetic inference analy- ses were conducted using Maximum Parsimony (MP), Bayesian inference (Bi) and Maximum Likelihood (ML), revealing that even a combined dataset of both mitochondrial and nuclear genes is not able to resolve the phylogenetic relationships of the Lacertidae family under the tribe level.
Karaman, S. (1921) -
Klemmer, K. (1957) -
Knauer, F. (1905) -
Knoblauch, A. (1907) -
Korsós, Z. (2002) -
Kramer, G. & Mertens, R. (1938) -
Kretzschmar, K.G. (1995) -
The dangers of helminth infestations for lizards, particularly infestations with nematodes und cestodes, are reported. Infection routes, diagnoses, und treatments are described.
Kroniger, M. & Zawadzki, M. (2005) -
Kwet, A. (2005) -
Kwet, A. (2006) -
Kwet, A. (2014) -
Kwet, A. (2017) -
Laforgia, V. & Capaldo, A. & Muoio, R. & Sciarrillo, R. (2001) -
Langerwerf, B. (1980) -
Lantermann, W. & Lantermann, Y. (2017) -
Lantz, L.A. (1927) -
Lapini, L. (2015) -
After a short synthesis of the present knowledge on the distribution of the Horvath’s rock lizard (Iberolacerta horvathi) in Italy, the Author refers about the discovery of a second population of the species on the Dolomites (Italia north-eastern Italy, Veneto Region). It dwells on the climbing wall of mount Tudaio (960 m a. s. l., Vigo di Cadore, Belluno) and has been detected thanks to the recovery of one Horvath’s rock lizard predated by a smooth snake female (Coronella austriaca) falling down from about 15 meters high. From its throat has been extracted a male of Iberolacerta horvathi determined by means of already known diagnostic characters: wide contact between rostral and frontonasal scales, flat and polygonal dorsal scales, caudal rings of regularly alternated thickness (one about a half of the other), bright white unspotted throat, bright yellow-lemon bellies. Status and consistency of the new population of Horvath’s rock lizard is not well known, but it is possible to note that at the bottom of the climbing wall the common wall lizard (Podarcis muralis) seems to be quite frequent.
Lapini, L. (2017) -
After a short synthesis of the present knowledge on the distribution of the Horvath’s rock lizard (Iberolacerta horvathi) in Italy, the Author refers about recent news and field verifications on the distribution of the lizard in Veneto and Friuli Venetia Giulia Regions (north-eastern Italy).
Lapini, L. & Dal Farra, A. (1994) -
Lapini, L. & Dall’Astra, A. & Luiselli, L. & Nardi, P. (2004) -
After a provisional review of the knowledge on Lacerto horvathi in Italy, the authors present new data on its distribution, spacing strategy and territoriality. The species is quite common in various mountainous habitats of north-eastern Italy, both in Friuli- Venezia Giulia (Udine Province, more than 55 localities; Pordenone Province, 1 locality) and Veneto (Belluno Province, 1 locality). In these areas, it lives from 250 (Pradolino Gorge, Pulfero, Udine) to 2000 metres a.s.l.. (M.te Ponza, Tarvisio, Udine). The preliminary comparison between the homing patterns of syntopic L. horvathi and Podarcis muralis suggests that this last-mentioned species is characterised by wider home ranges — 3-9 times bigger than those of L. horvathi — showing a very high inter-specific aggressiveness. However, in deeper gorges, in cool valleys with Dinaric orientation, as well as on top of mountains, L. horvathi tends to vicariate P. muralis for various differences in both spacing, thermal, and reproductive strategies. In north-eastern Italy, such vicariance on average gradually occurs between 700 and 900 meters above sea level.
Lapini, L. & Richard, J. (2006) -
Lapini, L. & Richard, J. & Dal Farra, A. (1993) -
Lescure, J. (2008) -
Lizana, M. & Carbonero, J. (2007) -
Lizana, M. & Martín-Sánchez, R. & Morales, J. & López-González, J. & Gutiérrez, J. (1993) -
Ljubisavljevic, K. (2019) -
Ljubisavljević, K. & Glasnović, P. & Kalan, K. & Kryštufek, B. (2012) -
In this paper, we present data on the female reproductive traits of the Horvath’s rock lizard from Slovenia. The clutch, egg and hatchling characteristics were investigated based on clutches laid in laboratory conditions by pregnant females collected from a natural population. A female lays one clutch of eggs annually with an average number of three (range 1-5) eggs. We found a significant positive correlation between female size and egg width and volume. The mean egg length and volume in a clutch decreased significantly with clutch size independently of female size. The incubation period averaged 44 days. Significant positive correlations were found between the hatchling total length and mass, and egg mass. There was a significant negative relationship between the egg mass and incubation duration. The life-history strategy of the Horvath’s rock lizard appeared to be shaped by several factors, such as the constraints of a high-altitude environment, flattened body morphology and possibly phylogenetic background.
Lončar, M. (2005) -
López, P. & Martin, J. (2002) -
Trade-offs between reproduction and survival are important determinants of life-history characteristics of lizards. Organisms cannot increase the allocation of limited resources to reproduction without diverting a proportional amount of energy from another trait. Locomotor performance is an ecologically relevant trait that potentially influences survival by affecting the ability to escape from predators. Most studies have used female lizards as subjects because pregnancy is known to reduce their locomotor abilities, whereas little is known on costs of reproduction in males. In this study we suggest that in males of the lizard Lacerta monticola reproductive investment in morphological traits that confer dominance (i.e. head size) might lead to a low probability of survival by decreasing investment in other traits that affect locomotor performance (i.e. limb symmetry). We staged laboratory agonistic encounters between males and measured their morphology and burst speed on a race track to examine possible relationships between morphology, social dominance and locomotor capacity. Our results indicate that social dominance was positively related to relative head height, and that escape speed was negatively related to levels of fluctuating asymmetry in femur length, but also negatively related to relative head height. Males with greater relative head height also had more asymmetrical femurs, thus dominant males suffered a decrease in locomotor performance. Males with higher heads tend to dominate male–male interactions and hence may gain access to reproductive females, thus increasing their current reproduction success. However, this might occur at the expense of future survivorship mediated by a decrease in escape speed. Therefore, in male L. monticola there might be a trade-off between current reproductive success and survival.
López, P. & Amo, L. & Martin, J. (2006) -
In spite of the importance of chemoreception in social organization and sexual selection of lizards, there is a lack of general knowledge on how the characteristics of chemical signals mediate these behaviors. Moreover, it is unknown which are the mechanisms that might confer honesty to the information provided by chemical signals. We analyzed here whether characteristics of the lipophilic fraction of femoral gland secretions of Lacerta monticola male lizards can be related to the morphology, physical condition, and health state of the sender. Our results indicated that some male traits, such as body size, number of blue spots, and number of femoral pores and their level of fluctuating asymmetry, were related to variability in the relative proportions of some lipophilic chemical compounds found in secretions. Thus, conspecifics could obtain reliable information on the producer of a scent mark based on chemicals alone, and this might be the basis of female choice observed in this lizard. Moreover, only males with a greater T-cell immune response had higher proportions of two steroids (ergosterol and dehydrocholesterol) in their femoral secretions, which might suggest that the signal is honest and costly to produce. We suggest that only high-quality males could divert these compounds from metabolism to secretions in order to produce an exaggerated and honest `chemical ornament.`
López, P. & Aragón, P. & Martin, J. (1998) -
López, P. & Aragón, P. & Martin, J. (2003) -
Female mate choice based on visual traits appears to be rare in lizards. Field observations suggest that females of the lizard Lacerta monticola preferred to mate with larger/older males. Although older males are usually green and larger, and younger males brown and smaller, there is some overlap in size and coloration between age classes. Thus, visual cues may not always be reliable indicators of a male`s age. We hypothesized that female mate-choice preferences may be based on males` pheromones, which might transmit information about characteristics such as age. In a laboratory experiment, we analyzed the effect of age of males on attractiveness of their scents to females. When we offered scents of two males of different age, females associated preferentially with scents of older males. This suggested that females were able to assess the age of males by chemical signals alone, and that females preferred to be in areas scent-marked by older males. Thus, females may increase their opportunities to mate with males of high quality, or may avoid harassment by sneaking young males. This result agreed with field observations on females mating with old males, and rejection of advances by young males. Our results also suggested that female preference for older males may depend on their own body size. Large females showed a strong preference for older males, whereas smaller females were not so selective. This, together with males` preference for large females, might lead to size-assortative matings. We suggest that the quality and/or quantity of male pheromones could communicate to the female heritable male genetic quality (i.e. age) and thereby serve as the basis of adaptive female choice in lizards.
López, P. & Hawlena, D. & Polo, V. & Amo, L. & Martin, J. (2005) -
Individual animals from the same population, sex, age and reproductive status often respond differently and consistently to predators. One important dimension of this behavioural variation is the shy–bold continuum. Innate differences in boldness might explain why individuals differ in their antipredator behaviour. In a laboratory experiment, we examined the sources of individual variation in antipredator behaviour of adult male lizards Lacerta monticola. We simulated in the laboratory repeated predatory attacks of low or high risk and analysed activity levels and refuge use in both situations. Multivariate analyses suggested the existence of two consistent and independent shy–bold continua. The first described a gradient from bold lizards that spent shorter times in the refuge after predatory approaches to shy lizards with longer emergence times, whereas the other described a gradient from bold lizards with a low propensity to hide when the predator was close but risk was low to shy lizards that hid more often. We analysed whether morphological characteristics, body condition and health (estimated from their T cell immunocompetence) of individuals might account for the differences observed. Bold individuals had smaller absolute body size, but relatively larger heads, better body condition and better health. Bold individuals with a low propensity to hide when risk was low had larger absolute body sizes, whereas relative head size, and body condition and health were not important. We suggest that the position of an individual in the shy–bold continua might reflect its optimal antipredator behaviour, which would be a function of its health, general quality and ability to evade predators.
López, P. & Luque-Larena, J.J. & Martin, J. (2003) -
López, P. & Martín, J. (2005) -
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.
López, P. & Martin, J. (2006) -
López, P. & Martín, J. (2012) -
Multiple sexual signals may convey information on males’ quality. However, most research focused on visual signals, ignoring chemical signals. In vertebrates, chemical signals are probably often a multicomponent mixture of several active compounds, but they are not well known. We examined the potential additive and interactive effects of 2 compounds (oleic acid and ergosterol) naturally found in femoral gland secretions of male rock lizards on chemosensory exploration behavior of females. Tongue-flick (TF) rates of females to male secretions may result from the combination of responses to multiple compounds. There may be an additive or synergetic effect because male secretions with the highest proportions of both compounds received the highest TF rates, suggesting that their scents were more intriguing. However, there might be an interactive effect too; female TF rates were higher to males’ scent with high proportions of ergosterol alone, even if proportions of oleic acid were low, than to high proportions of oleic acid but with low proportions of ergosterol. Further bioassays testing TF behavior of females to standard compounds, presented alone or combined in different concentrations confirmed these findings. Variations in female TF behavior might be explained because different compounds signal different male traits of different importance for females. Our study suggested that femoral secretion of male rock lizards may act as a multicomponent chemical signal.
López, P. & Martin, J. & Cuadrado, M. (2004) -
Vertebrate males often show breeding colours that may function as reliable signals of status in intrasexual competition. In many lacertid lizards, males show a conspicuous row of small but distinctive blue spots that runs along their body side on the outer margin of the belly. However, no study has examined the role of these blue spots. We first analysed in a field population of the Iberian rock lizard, Lacerta monticola, the relationships between number of blue spots and some morphological traits, which are known to be related to males’ fighting ability. The number of spots seems to be an character showing ontogenetic change as large (generally older) males showed more blue spots than small (generally younger) males. Males with a higher body condition also showed a higher number of blue spots. Thus, a higher number of blue spots may be used to signal size, age or body condition. Many contiguous blue spots would result in a visual artefact consisting of a continuous blue band, which might be a reliable size- or condition-dependent signal in some social contexts. We further examined in the laboratory whether male characteristics are related to dominance status. In males with similar body size or age, those with relatively larger heads were more dominant, whereas the number of blue spots was not important. Moreover, the number of blue spots in nature was not related to relative head size. Finally, we experimentally manipulated the presence and the number of blue spots of intruding males, and examined the aggressive response of resident males. Intruder individuals manipulated to cover all their blue spots received a lower amount of aggression. However, males with different numbers of manipulated blue spots received a similar number of aggressive responses. These results suggest that, during agonistic encounters, the presence of blue spots, but not their number, may elicit aggressiveness. Thus, blue spots may serve to identify an individual as an adult male, and to enhance body size of larger males.
López, P. & Moreira, P.L. & Martin, J. (2009) -
In the lizard Iberolacerta monticola, there are 2 discrete color morphs (`blue` vs. `green`) that seem to have alternative reproductive strategies. Because scent marking of territories and chemoreception are important in social organization of these lizards, we explored whether there is also chemical polymorphism and chemosensory recognition between color morphs. Analyses by gas chromatography-mass spectrometry showed that adult males of different morphs had similar chemical compounds in femoral gland secretions, but the proportions of some shared chemicals were different; blue morph males had higher proportions of steroids and lower proportions of fatty acids than green males. Differential tongue-flick rates to scents from femoral secretions of males indicated that males were able to detect the scent of any male from a baseline odorless control (water) but did not discriminate between the scent of males of different morphs. In contrast, females detected and discriminated between scents of the 2 male morphs by chemical cues alone. We suggest that differences in males` chemical signals between morphs and the ability of females of discriminating chemically between male morphs might be required to maintain spatial behavior, social organization, and mate choice decisions and, thus, be important for the maintenance of alternative reproductive strategies and polymorphism in lizards.
López, P. & Muñoz, A. & Martin, J. (2002) -
Female preference for dominant males is widespread and it is generally assumed that success in male-male competition reflects high quality. However, male dominance is not always attractive to females. Alternatively, relatively symmetric individuals may experience fitness advantages, but it remains to be determined whether males with more symmetrical secondary sexual traits experience advantages in both intra- and intersexual selection. We analysed the factors that determine dominance status in males of the lizard Lacerta monticola, and their relationship to female mate preference, estimated by the attractiveness of males` scents to females. Sexually dimorphic traits of this lizard (head size and femoral pores) appear to be advanced by different selection pressures. Males with relatively higher heads, which give them advantage in intrasexual contests, were more dominant. However, head size was unimportant to females, which preferred to be in areas marked by relatively heavier males, but also by males more symmetric in their counts of left and right femoral pores. Chemicals arising from the femoral pores and other glands might honestly indicate quality (i.e. related to the symmetry levels) of a male to females and may result from intersexual selection. Females may use this information because the only benefit of mate choice to female lizards may be genetic quality. Chemical signals may be more reliable and have a greater importance in sexual selection processes of lizards than has previously been considered.
Loureiro, A. & Almeida, N.F. de & Carretero, M.A. & Paulo, O.S. (2008) -
Loureiro, A. & Ferrand de Almeida, N. & Carretero, M.A. & Paulo, O.S. (2008) -
Luca, N. de (1989) -
External morphology of Horvath`s rock lizard (Lacerta liorvathi MEHELY 1904) was examined. Variability was established in the biornetric, meristic and qualitative characters, as well as differences between the adult and juvenile an males and females and between the samples of animals from the Slovenian and north Dinarie part of the area, New localities were stated and the known area of this species expanded.
Luca, N. de (1990) -
Luca, N. de (1992) -
Lacerta horvathi is a relatively poorly known species assigned to a taxonomi- cally problematic group of lacertid lizards - Archaeolacerta. It is present with scattered mountain populations in the Julian Alps of Slovenia, in the Ucka Moun- tain and the northern Dinarid Mountains of Croatia, in northeastern Italy, in southwestern Austria, and in southern Germany (LAPINJ & DOLCE 1983, GRIL- LITSCH & TEDEMANN 1986, DE LUCA 1989, CAPULA & LUISELLI 1990). Much of data on distribution and taxonomy of this species were presented during the past few years (e.g. the papers cited above, DE LUCA & DULIC 1988, CAPULA et al. 1989). However, the comprehensive data regarding biology and ecology of L. horvathi are still missing. In this paper some new data on daily and seasonal activity patterns, thermoregulation, reproduction and food composition, are presented.
Luca, N. de & Đulić, B. (1988) -
Karyotype analysis of 2 male horvath`s rock lizard, Lacerta (Archaeolacerta) horvathi, found 2n = 36 in both somatic and spermatogonial metaphases. All the chromosomes are acrocentric. In diakinesis, 18 bivalents are visible. Microchromosomes, characteristic of most lacertid karyotypes, are absent in this species. In a single metaphase, satellites were observed on the 8th pair of chromosomes. Male sex chromosomes have not been determined. Overall, the karyotype of Lacerta horvathi is similar to that of Lacerta vivipara.
Luca, N. de & Kleteèki, E. & Dulic, B. (1989) -
Lutz, D. & Mayer, W. (1985) -
Albumins from several species assigned to Lacerta s. 1. and Gallotia were investigated by means of the Micro-Complement fixation technique. The phylogenetic trees evaluated according to Fitch & Margoliash (1967) and Beverley & Wilson (1982) led us to infer a division, which is in accordance with protein electrophoretic results (Mayer & Tiedemann, 1982) as well as with the subgeneric division used before the studies of Böhme (1971) and Arnold (1973).
Malkmus, R. (1981) -
ln der vorliegenden Arbeit wird die Herpetofauna von über 50 portugiesischen Gebirgszügen zusammengestellt: für jedes Gebirge der hõchste bisher bekannte Fundort der einzelnen Arten mit Hôhenangabe. Es ist das Ergebnis vierjiähriger Untersuchungen (1976-1980).
Malkmus, R. (1982) -
This survey is presenting the geographical range of 17 amphibian and 29 reptile species (including the marine turtles) known to occur in Portugal, based on available data from the literature as well as ehe author`s own observations during the years 1976- 1981. It is the first attempt to map exactly the herpetofauna of a southern European country. Data are given concerning the horizontal and vertical distribution of each species, the density of distribution as well as the typical habitats, the latter being described in some detail. The rhythm of activity of most of the species depends on the different climatic conditions within this country. The few available phenological data are sum- marized.
Malkmus, R. (2013) -
Mallard, F. (2018) -
Understanding the local effects of climate change on biodiversity is essential to guide environmental and management policies for natural areas. The lack of knowledge at the regional level has led to the development of a research program `Climate sentinels`. The main hypothesis of the program is that effects on biodiversity are particularly detectable in species that have low movement abilities. These `climate sentinel` species would be the first to respond to local climatic variations by adaptation or local extinction. From the territory of New Aquitaine (France), 20 biological indicators of climate change in different ecosystems (dune, dry, wet, mountain and forest) were chosen: plants communities of ecosystems, insects (butterflies, dragonflies, bumblebees), amphibians (Rana pyrenaica, Hyla molleri, H. arborea), reptiles (Iberolacerta bonnali, Timon lepidus, Zootoca vivipara, Emys orbicularis, Vipera berus) and mammals (Marmota marmota, micromammals). This paper presents the method of development of these indicators and the research hypotheses associated with climate sentinel species.
La compréhension des effets locaux du changement climatique sur la biodiversité est essentielle pour orienter les politiques environnementales et de gestion des espaces naturels. Le manque de connaissances au niveau régional a conduit au développement d’un programme de recherche « les sentinelles du climat ». L’hypothèse principale du programme est que les effets sur la biodiversité sont particulièrement détectables chez des espèces qui ont de faibles capacités de déplacements. Ces espèces « sentinelles du climat » seraient les premières à répondre aux variations climatiques locales par adaptation ou par extinction locale. À partir du territoire de la Nouvelle-Aquitaine, 20 indicateurs biologiques du changement climatique dans différents écosystèmes (dunaire, sec, humide, montagnard et forestier) ont été choisis : cortèges floristiques des écosystèmes, insectes (papillons, libellules, bourdons), amphibiens (Rana pyrenaica, Hyla molleri,H. arborea), reptiles (Iberolacerta bonnali, Timon lepidus, Zootoca vivipara, Emys orbicularis, Vipera berus) et mammifères (Marmota marmota, micromammifères). Cet article présente la méthode de développement de ces indicateurs et les hypothèses de recherche associées aux espèces sentinelles du climat.
Mangiacotti, M. & Martin, J. & López, P. & Reyes-Olivares, C.V. & Rodriguez-Ruiz, G. & Coladonato, A.J. & Scali, S. & Zuffi, M.A.L. & Sacchi, R. (2020) -
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.
Marchi, G. de & Bombieri, G. & Boz, B. & Leandri, F. & Richard, J. (2020) -
The Horvath’s rock lizard (Iberolacerta horvathi) is a rupicolous mountain species endemic of the eastern Alps and northern Dinaric range. The species has its known western limit of the distribution in the Veneto region of Italy. It is not known whether the species is really rare in Veneto or whether the area has been insufficiently surveyed. In addition, it is not known whether the westward distribution of the species is limited by a physiographic or by a climatic barrier. During the period 2016-2018, 118 sites were surveyed in the Veneto and Trentino-Alto Adige regions. Four new occurrences of Iberolacerta horvathi were discovered in Veneto that: 1) largely fill the gap between the westernmost known site and the closest site to the east; 2) extend further west the known distribution by 9 km. In addition the species was confirmed in three already known sites. A species distribution model was developed with the software MaxEnt, using 100 occurrences from Italy, Austria and Slovenia. The best model shows that the distribution is explained by the asperity of their habitat, the sedimentary bedrock, the aspect, the average temperature of the coldest quarter, the rainfall seasonality and the average summer rainfall. The last variable appears as the most likely responsible for the rarefaction of the species at its western limit. In addition, the species distribution model suggest that the Horvath’s rock lizard might be present in some additional mountain groups where it has so far not been found yet
Marchi, G. de & Boz, B. & Bombieri, G. & Richard, J. (2019) -
Marco, A. & López-Vicente, M. & Pérez-Mellado, V. (2004) -
Martín, J. (2001) -
Prey often respond to predators by increasing their use of refuges but relatively few studies have analyzed how prey decide when to resume their behavior after a predator’s unsuccessful attack. This is important because refuge use may have some costs that should be minimized such as the loss of time available for foraging or mate searching. In addition, unfavorable conditions in refuges (e.g., suboptimal temperatures) might entail physiological costs such as hypothermia. Under these circumstances, animals should optimize the decision of when to come out from a refuge by balancing the fitness effects of the diminution of predation risk with time against the costs of loss of time available for other activities and loss of time spent at optimal body temperature. I review several experiments with lizards that support that individuals decide to come out from a refuge when the costs of hiding exceed predation risk in the exterior, and that there is an optimal emergence time. Optimization of these antipredatory behavioral strategies might help lizards to cope with changes in predation risk without incurring excessive costs.
Martín, J. (2009) -
Martin, J. (2009) -
Martín, J. & López, P. (1999) -
Prey often respond to predator presence by increasing their use of refuges. However, unfavorable thermal conditions in refuges might entail physiological costs for an ectothermic prey. Thus, the decision of when to come out from a refuge should be optimized by considering the expected fitness effects of diminution of predation risk with time, but also by considering the cost of the loss of time spent at optimal body temperature maximizing physiological functions. The model of Ydenberg and Dill describes the trade-off between risk and cost for a prey fleeing to a refuge. We present a special case of this model to predict how emergence time from the refuge in lizards or other ectotherms should vary as a function of risk of predation and thermal costs of refuge use. The analyses of the variation in emergence time from a refuge of Lacerta monticola lizards in the field under two different predation risk levels supported the predictions of the model. As predicted, time spent in the refuge was longer when the threat of the initial attack had been higher, and therefore the subsequent diminution of risk was slower, but only when lizards emerged at the same place where they hid. When initial body temperature was high, some lizards decreased emergence time by emerging from a different place. In addition, the effects of thermal costs were more relevant in the high-risk situation. Time spent in the refuge under high risk increased when thermal conditions of the refuge were more similar to thermal conditions outside (i.e., physiological costs of refuge use were lower). We conclude that optimization of refuge-use strategies might help lizards cope with changes in predation risk without incurring excessive physiological costs.
Martin, J. & López, P. (2000) -
Theoretical models of anti-predator escape behaviour suggest that prey may adjust their escape response such that the optimal flight distance is the point at which the costs of staying exceed the costs of fleeing. Anti-predatory decisions should be made based also on consequences for long-term expected fitness, such as the costs of refuge use. For example, in lizards, the maintenance of an optimal body temperature is essential to maximize physiological processes. However, if unfavourable thermal conditions of refuges can decrease the body temperature of lizards, their escape decision should be influenced by refuge conditions. Analyses of the variation in flight distances and emergence latency from a refuge for the lizard Lacerta monticola under two different predation risk levels, and their relationship with the thermal environment, supported these predictions. When risk increased, lizards had longer emergence latencies, and thus costs of refuge use increased (a greater loss of time and body temperature). In the low-risk situation, lizards that were farther from the refuge had longer flight distances, whereas thermal conditions were less important. When risk increased, lizards had longer flight distances when refuges were farther off, but also when the external heating rate and the refuge cooling rate were lower. The results suggest that, in addition to the risk of predation, expected long-term fitness costs of refuges can also affect escape decisions.
Not all individual lizards in a population are simultaneously active even when thermal conditions are suitable for activity. We examined inter-individual variability in activity levels of male Iberian rock lizards (Lacerta monticola) in a seminatural enclosure during the mating season, and analyzed whether social status affects their activity levels, time budgets, and body-mass changes. Activity levels of lizards varied significantly with time of day. However, activity levels of individual males were significantly correlated with their rank in the social hierarchy. When the males were active, their status did not influence the time spent basking, resting, or moving, but males with a higher status spent more time in social activities. Higher activity levels were costly, causing males to lose more body mass, although this could have been mainly due to the costs of maintaining a higher social status. We conclude that because attaining a higher status may require a male to be more active and more involved in agonistic encounters, subordinate individuals decrease their activity in order to decrease the costs of social behavior.
Martin, J. & López, P. (2001) -
Prey often respond to predator presence by increasing their use of refuges. However, because the use of refuges may entail several costs, the decision of when to come out from a refuge should be optimized. In some circumstances, if predators remain waiting outside the refuge and try new attacks or if predator density increases, the prey may suffer successive repeated attacks in a short time. Successive attacks may represent an increase in the risk of predation, but the costs of refuge use also may increase with time spent in the refuge. Thus, prey should make multiple related decisions on when to emerge from the refuge after each new attack. We simulated in the field repeated predatory attacks to the same individuals of the lizard Lacerta monticola and specifically examined the variation in successive times to emergence from a refuge under different thermal conditions (i.e., different costs of refuge use). The results showed that risk of predation but also thermal costs of refuge use affected the emergence decisions. Lizards increased progressively the duration of time spent in the refuge between successive emergence times when the costs of refuge use were lower, but tended to maintain or to decrease the duration of time spent in the refuge between successive emergence times when cost of refuge use increased. Additionally, lizards that entered the refuge with higher body temperatures had overall emergence times of longer duration. Optimization of refuge use and flexibility in the antipredator responses might help lizards to cope with increased predation risk without incurring excessive costs of refuge use.
Martin, J. & López, P. (2003) -
In lizards, ontogenetic changes in body size affect thermal-exchange rates. This simple physical property may have consequences for thermoregulation, and also for antipredator behavior. We examined how ontogenetic changes in body mass affect rates of heating and cooling of the lizard Lacerta monticola, confirming the general result obtained for other lizards. We further analyzed the differences between juveniles and adults in approach distances to a simulated predator and in time to emerge from refuges. Juvenile lizards have a lower absolute running speed, making them more vulnerable to predation. However, in contrast to results expected from optimal-escape theory, approach distances were shorter for juveniles than for adults. Juveniles may be confident in their small size and only flee when the probability of being detected is high. On the other hand, differences in thermal properties might affect costs of refuge use. Thus, juveniles might delay fleeing because their costs of hiding are higher, as they cool faster than adults. Differences in thermal costs may also explain the juveniles` shorter times of emergence from refuges. Because of the behavioral adjustments involved in antipredator behavior, the physiological costs of reaching a low body temperature in refuges probably do not differ between age classes.
Prey often respond to predator presence by increasing their use of refuges, but because this strategy may be costly, the decision regarding when to come out from a refuge should be optimized. The loss of foraging opportunities may be one of the main costs when safer microhabitats (i.e. refuges) are also the poorest in terms of their foraging profitability. We present the results of an experimental field study to test whether emergence times from a refuge of the Iberian rock lizard, Lacerta monticola, vary as a function of expected foraging opportunities and level of satiation of the lizard. As predicted, short-term fluctuations in availability of food influenced emergence times; when a lizard had just detected some food in the recent past, emergence times decreased greatly, because the loss of opportunities for foraging increased costs of refuge use. Furthermore, the characteristics and success of the encounter with food, nutritional state of lizards, and the added possibility of capturing new food items influenced the duration of hiding times. Therefore, foraging requirements and avoidance of predators may be conflicting demands that L. monticola lizards balance by modifying the duration of time spent in refuges.
Martin, J. & López, P. (2006) -
In spite of the fact that chemoreception plays an important part in sexual selection of many vertebrates, there is a lack of general knowledge on the role of specific chemical compounds as sexual signals, and on how the characteristics of chemical signals mediate behaviours such as mate choice.
We examined how female Iberian Rock Lizards use the chemical characteristics of femoral gland secretions of males to select potential mates.
Tongue-flick assays and choice of scent trials showed that females discriminated by chemosensory cues alone differences in chemical secretions of different males. Females showed stronger chemosensory responses and preferred the scent of males of presumably high quality (i.e. those more symmetric and with a greater immune response).
Chemical analyses by gas chromatography-mass spectrometry showed that these preferred males allocated relatively more cholesta-5,7-dien-3-ol and ergosterol to femoral secretions, which suggested that females used these chemicals to choose between males’ scents.
Further experiments confirmed that females were able to discriminate cholesta-5,7-dien-3-ol, ergosterol, and changes in their concentrations, from similar steroids (i.e. cholesterol) also found in secretions of males. Moreover, females were more attracted to areas that were experimentally manipulated to increase the proportion of ergosterol in natural scent marks of males.
We suggest that femoral secretions with higher proportions of cholesta-5,7-dien-3-ol and ergosterol might be reliable advertisements of quality of a male, which females could use to select mates.
Evolutionary theory proposes that signals used in sexual selection can only be stable if they are honest and condition dependent. However, despite the fact that chemical signals are used by many animals, empirical research has mainly focused on visual and acoustic signals. Vitamin D is an essential nutrient for lizards, but in some lizards its precursor (cholesta-5,7-dien-3-ol=provitamin D) is found in femoral gland secretions, which males use for scent marking and intraspecific communication. By allocating provitamin D to secretions, males might need to divert vitamin D from metabolism. This might be costly and condition dependent. We tested whether diet quality affected chemical signals of male Iberian rock lizards (Lacerta monticola) and its consequences for sexual selection. After experimental supplementation of dietary vitamin D, males increased the proportion of provitamin D in femoral secretions. Further experiments showed that females detected these changes in males` signals by chemosensory cues, and discriminated provitamin D, and changes in its concentration, from similar steroids (i.e. cholesterol) found in secretions. Moreover, females preferred areas scent marked by males with more provitamin D in their secretions. This mechanism would confer honesty to chemical signals of male lizards, and, thus, females may rely on it to select high-quality males. We suggest that the allocation of vitamins and other essential nutrients to either visual (e.g. carotenoids) or chemical ornaments might be the common basis of honest sexual displays in many animals.
Martin, J. & López, P. (2007) -
Intrasexual competition favours the evolution of conspicuous fighting ability badges. However, in spite of the fact that chemoreception is important in sexual selection of many animals, such as lizards, the role of chemical signals in males` contests is relatively unknown. Here, we show that proportions of cholesterol in femoral gland secretions of male Iberian rock lizards were related to their body size (which confers a competitive advantage in fights). Males discriminated chemically and responded aggressively to cholesterol stimuli presented on swabs. Moreover, we experimentally increased cholesterol in the scent of males, and staged encounters in neutral cages between two unfamiliar and size-matched males. Focal males lost more agonisitic interactions against males manipulated with cholesterol than in control tests. We suggest that differences in scent composition may reliably signal fighting ability in many lizard species, which would help to avoid the costs of fighting.
Martin, J. & López, P. (2008) -
Some mate choice theories propose that only male signals that are honest and condition-dependent can be stable, while another hypothesis states that males evolve signals that exploit the sensory system of females. However, sensory traps might evolve into honest signals if they are differentially costly for males. We tested whether a pre-existing sensory bias for food chemicals explained chemosensory preferences of female Iberian rock lizards for male scents. We manipulated hunger levels of females and found that food-deprived females had increased chemosensory responses to chemical stimuli from both invertebrate prey and femoral secretions of males, but not to control water. Further tests suggested that cholesta-5,7-dien-3-ol (provitamin D3), a lipid found in both prey and males’ scent, may be one of the chemicals eliciting these responses. Moreover, hungry females spent more time on scent marks of males that had experimentally increased cholesta-5,7-dien-3-ol than on scent marks of males alone, whereas for control females this effect was not significant. We suggest that preexisting sensory bias for essential nutrients (i.e., provitamin D) may be the origin of similar female responses to male chemicals. However, previous studies have suggested that the allocation of these chemicals to ornaments is costly and only high quality males can afford it. Therefore, preexisting sensory bias for essential nutrients may further allow the evolution and maintenance of honest sexual display.
Martín, J. & López, P. (2010) -
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.
Martin, J. & López, P. (2012) -
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.
Martín, J. & López, P. (2013) -
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.
Martín, J. & López, P. (2014) -
Many lizards have diverse glands that produce chemosignals used in intraspecific communication and that can
have reproductive consequences. For example, information in chemosignals of male lizards can be used in
intrasexual competition to identify and assess the fighting potential or dominance status of rival males either
indirectly through territorial scent-marks or during agonistic encounters. Moreover, females of several lizard
species “prefer” to establish or spend more time on areas scent-marked by males with compounds signaling a
better health or body condition or a higher genetic compatibility,which can have consequences for theirmating
success and inter-sexual selection processes. We review here recent studies that suggest that the information
content of chemosignals of lizards may be reliable because several physiological and endocrine processes
would regulate the proportions of chemical compounds available for gland secretions. Because chemosignals
are produced by the organismor come fromthe diet, they should reflect physiological changes, such as different
hormonal levels (e.g. testosterone or corticosterone) or different health states (e.g. parasitic infections, immune
response), and reflect the quality of the diet of an individual. More importantly, some compounds thatmay function
as chemosignals also have other important functions in the organism (e.g. as antioxidants or regulating the immune
system), so there could be trade-offs between allocating these compounds to attending physiological needs or to
produce costly sexual “chemical ornaments”. All these factors may contribute to maintain chemosignals as
condition-dependent sexual signals, which can inform conspecifics on the characteristics and state of the sender
and allow making behavioral decisions with reproductive consequences. To understand the evolution of chemical
secretions of lizards as sexual signals and their relevance in reproduction, future studies should examine what
information the signals are carrying, the physiological processes that can maintain the reliability of the message
and how diverse behavioral responses to chemosignals may influence reproductive success.
Martín, J. & López, P. & Cooper, W.E. (2003) -
Because time spent in refuge may be costly if prey lose opportunities to forage, fight, or mate, prey allow predators to approach closer before beginning to flee when opportunity costs are high. Because the same opportunity costs may apply to refuge use as to escape, prey should make similar trade-offs between risk of emerging and cost of remaining in refuge. In the Iberian rock lizard, Lacerta monticola, we studied the effects of sex, reproductive season, speed of predator approach, and potential loss of mating opportunities on time spent in refuge following simulated predatory attacks. Lizards of both sexes adjusted refuge use to the level of risk by spending more time in refuge when approached rapidly than slowly. Females remained in refuge for equal times in the mating and postreproductive seasons, but males emerged sooner during the mating season, suggesting adjustment to a cost of lost opportunity to search for mates during the mating season. When a tethered female was nearby, males emerged from refuge earlier than if no female was present, indicating a trade-off between risk and mating opportunity. Approach speed affected emergence time when females were absent, but not when a female was present. Approach speed did not affect the probability that, after emerging, a male would return to court the female. For males that courted females intensely (bit them) before entering refuge, approach speed did not affect latency to emerge, but males that courted less intensely emerged sooner if approached slowly than rapidly. These findings show that males adjust the length of time spent in refuge to both risk of predation and reproductive cost of refuge use.
Martin, J. & Lopez, P. & Carrascal, L.M. & Salvador, A. (1995) -
We studied the relationships between thermoregulatory basking postures and heating rates in the Iberian rock lizard (Lacerta monticola). Heating rates were high when body size was small, the substrate temperature was high, and the angle of incidence of the sun`s rays was high (i.e., the rays were perpendicular to the lizard`s body). However, heating rates were not related to the compass orientation of lizard`s body axis to the sun (perpendicular versus in line with the sun). In the field, the slopes of the body axis of basking individuals were higher during the early morning. However, neither the cardinal orientation of a lizard`s body axis to the sun nor the frequency of flattening varied during the morning. Postural adjustments to select an adequate angle of incidence of the sun`s rays on the dorsal part of the body suggest that this behaviour might enhance absorption of solar radiation.
Martin, J. & López, P. & Polo, V. (2009) -
Prey experience a broad range of temporal patterns of predation risk and may face different problems when assessing risk level and deciding how to allocate costly antipredator behaviour across time. We simulated in the laboratory repeated predatory attacks of different intensity (low versus high risk) to Iberian rock lizards, Iberolacerta cyreni, with similar numbers of low- and high-risk attacks across a trial, but with different temporal patterns of presentation (i.e. a series of low-risk and then a series of high risk, or vice versa, or successive attacks where risk changed randomly). We measured activity levels, time hidden in refuges after each predatory attack, and propensity to hide in refuges when the predator is close but not actually attacking (‘false alarm’ flights). Antipredator responses were of higher intensity when risk was high, but responses also depended on temporal patterns of risk. When previous attacks were of low-risk or when risk was random, prey activity did not change between risk levels. False alarm flights in the low-risk situation were more frequent when risk was random or immediately after a series of high-risk attacks. When successive attacks were of similar risk level, lizards seemed able to predict risk level in the next attack and modified their antipredator behaviour accurately. However, when risk level was random, lizards did not seem able to predict risk level and adopted a conservative strategy, overestimating risk even though costs of the antipredator responses increased.
Martín, J. & Luque-Larena, J.J. & López, P. (2009) -
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.
Martín, J. & Moreira, P.L. & López, P. (2007) -
Male competition for females often results in the evolution of conspicuous male
traits that signal ﬁghting ability or dominance status. Most studies examining mechanisms
allowing these traits to function as reliable status signals (or badges) have focused on
conspicuous visual or acoustic traits. However, many animals communicate chemically,
and chemical traits alone may also signal male dominance status.
We examined whether chemicals in femoral gland secretions of male Iberian rock
Lacerta monticola monticola
Boulenger 1905) may signal dominance status.
In staged encounters, larger males were dominant over smaller ones. While controlling
for male body size, males of higher dominance status produced femoral secretions with
higher proportions of hexadecanol and octadecanol.
Tongue-ﬂick assays showed that males were able to (1) discriminate, by chemosensory
cues alone, the different concentrations of hexadecanol from other chemicals presented
in femoral secretions; (2) respond aggressively towards hexadecanol while responding
neutrally towards other chemicals; and (3) show differential chemosensory and aggressive
behaviours towards hexadecanol according to their own dominance status.
These results suggest that hexadecanol may be a reliable status badge. Moreover,
because hexadecanol elicits male aggressive behaviour, subordinate males signalling
high status (‘cheaters’) may end up paying high ﬁghting costs. In addition, males that
had higher dominance status, and that allocated higher proportions of hexadecanol to
femoral secretions, had greater T-cell immune responses. This result suggests a possible
link between quality of the immune system, dominance status, and chemical signals.
Martin, J. & Salvador, A. (1992) -
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.
Martin, J. & Salvador, A. (1993) -
The consequences of tail loss on the thermal biology of high-altitude rock lizards (Lacerta monticola) were studied by comparing field body temperatures, selected body temperatures in a gradient, and behavioural patterns of thermoregulation of tailed and tailless lizards in the field. Neither field nor selected body temperatures differed significantly between these groups. Time spent basking, mean duration of basking, and basking frequency did not differ between tailed and tailless lizards. Tailless lizards seemingly adjusted for running impairment by using rocks to a greater extent than did tailed lizards, and low morning body temperatures by increasing maximal duration of basking periods and keeping shorter distances to the nearest refuge. Tailless lizards modify thermoregulatory behaviour, but obtain the same thermoregulatory precision as tailed lizards, possibly allowing maximization of physiological processes linked to tail regeneration.
Many lizards autotomize their tail to escape when attacked by a predator. The foraging tactics of tailless lizards may be affected by changes in behavior aimed at compensating tail loss per se as well as associated costs of regeneration. We tested this hypothesis in a population of the Iberian rock-lizard (Lacerta monticola), during the reproductive season, by comparing the diet of tailed and tailless lizards with the estimated prey availability in their habitat. We also compared the feeding rates, foraging microhabitat and foraging tactics of tailed and tailless lizards. Tail loss did not affect feeding rates, nor did it alter the foraging tactics of either sex during spring. Tailless lizards used rocky microhabitats where predation was avoided, but where food availability was lower, and thus diet diversity decreased at least among tailless males. Their diet consisted primarily of Bibio flies, an apparently suboptimal prey item.
Many lizards autotomize their tails to escape when grasped by a predator. It is hypothesized that tail loss causes a reduction in social status, thereby potentially lowering their reproductive success. We experimentally induced tail loss in Lacerta monticola in a semi-natural enclosure, and show that tail loss reduced social status and mating access in males. Tailless males increased body mass more rapidly than tailed dominant males, probably due to lower aggression costs. Also, tailless females were courted less and copulated less than tailed females, supporting the hypothesis that tail loss decreases reproduction potential.
Martin, J. & Salvador, A. (1995) -
Martin, J. & Salvador, A. (1997) -
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.
Many lizards use caudal autotomy as an antipredatory strategy. We experimentally examined the effect of tail loss on the time-budgets, movement patterns, and home range size of the Iberian rock lizard Lacerta monticola in the field during the mating season. Our results indicate that tail loss did not alter most variables in male and female time-budgets but did affect the time spent moving and movement patterns of males. Tailless males spent less time moving and, when moving, more time on rocks, with shorter movements, more pauses, and less time on bushes than did tailed ones. Home range size was smaller in tailless males but did not differ between tailed and tailless females. The results suggest that tailless males may compensate the survival cost of autotomy by modifying their use of space and time; as a result, however, they may incur lower access to females.
Martinez-Rica, J.-P. & Reine-Vinales, A. (1988) -
Martinez-Rica, J.P. (1976) -
Martinez-Rica, J.P. (1977) -
Martinez-Solano, I. (2001) -
Martriun Rueda, J. (1992) -
Masó, A. & Pijoan, M. (2011) -
Matos, A. & Alves, P. (2001) -
Matos, A. de (1988) -
Mayer, W. (2013) -
A commented lacertids list for Europe, Africa, the Middle East, including the Arabian Peninsula and Asia is given.
Mayer, W. & Arribas, O. (2003) -
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.
Mayer, W. & Arribas, O.J. (1996) -
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.
Mayer, W. & Benyr, G. (1994) -
The albumins from 41 species of the family Lacertidae representing a substantial part of genera as well as
infrageneric groups of the collective genus Lacerta s.l. were investigated by means of the MCF technique.
The data allow a chronological estimation of the most important radiation events and their correlation with
paleogeographical facts. Our results are in remarkable contradiction to the phylogeny presented by ARNOLD
(1989a) based mainly on morphological characters. The most important phylogenetic relationships are:
1. The genera Gallona and Psammodromus are already isolated from the other taxa since the Oligocene.
We assess these main branches as subfamilies Gallotiinae and Lacertinae.
2. Lacerta s.str. and Lacerta subgenus Zootoca form the sister group of the remaining Lacertinae.
3. A colonization of Africa in the Lower Miocene has led to a separation in two lines, with mainly Eurasian
and African members respectively.
4. ARNOLD`S (I.e.) `Ethiopian and advanced Saharo-Eurasian clade` proved to be a paraphyletic unit, one
group of genera being more closely related to European taxa.
All techniques applied to the systematics of Lacertidae hitherto are critically discussed. An area-time-
hypothesis concerning the phylogeny of Lacertidae is presented.
Key words: Lacertidae, phylogeny of; Gallotiinae subfam. nov.; Lacertinae subfam. nov.; micro
complement fixation; albumin evolution.
Mayer, W. & Pavlicev, M. (2007) -
The family Lacertidae encompasses more than 250 species distributed in the Palearctis, Ethiopis and Orientalis. Lacertids have been
subjected in the past to several morphological and molecular studies to establish their phylogeny. However, the problems of convergent
adaptation in morphology and of excessively variable molecular markers have hampered the establishment of well supported deeper phylogenetic
relationships. Particularly the adaptations to xeric environments have often been used to establish a scenario for the origin and
radiation of major lineages within lacertids. Here we present a molecular phylogenetic study based on two nuclear marker genes and
representatives of 37 lacertid genera and distinct species groups (as in the case of the collective genus Lacerta). Roughly 1600 bp of
the nuclear rag1 and c-mos genes were sequenced and analyzed. While the results provide good support to the hitherto suggested main
subfamilies of Gallotiinae (Gallotia and Psammodromus), Eremiainae and Lacertinae [Harris, D.J., Arnold, E.N., Thomas, R.H., 1998.
Relationships of lacertid lizards (Reptilia: Lacertidae) estimated from mitochondrial DNA sequences and morphology. Proc. R. Soc.
Lond. B 265, 1939–1948], they also suggest unexpected relationships. In particular, the oriental genus Takydromus, previously considered
the sister-group to the three subfamilies, is nested within Lacertinae. Moreover, the genera within the Eremiainae are further divided into
two groups, roughly corresponding to their respective geographical distributions in the Ethiopian and the Saharo-Eurasian ranges. The
results support an independent origin of adaptations to xeric conditions in different subfamilies. The relationships within the subfamily
Lacertinae could not be resolved with the markers used. The species groups of the collective genus Lacerta show a bush-like topology in
the inferred Bayesian tree, suggesting rapid radiation. The composition of the subfamilies Eremiainae and Lacertinae as well as their
phylogeography are discussed.
Mazzotti, S. & Miserocchi, D. (2009) -
The collection of Amphibians and Reptiles of the Museum of Natural History of Ferrara has characteristics of historical representation, taxonomic and zoogeography that may be considered as examples of medium-sized types of collections more preva- lent in Italian museums. It consists of a historic part of the study and a donation from Guido Campadelli. The collection consists of 1739 specimens of amphibians (1734 of which held in 437 containers with ethanol and 5 taxidermized specimens) for a total of 78 amphibian species. There are 811 specimens of Reptiles (769 of which are stored in 526 containers with ethanol and 42 taxidermized specimens) representing a total of 108 species. The findings that form the collection of Amphibians come from 153 loca- lities of 15 states and 12 regions and 29 provinces of Italy. In the batrachological col- lection 62.1% of the total European species, and 94.3% of the total species in Italy, are represented. Zoogeographic regions most represented are the Palaearctic with 67% of the species of Amphibians in the collection and the Neotropical with 29%, fol- lowed by the Ethiopian (6%) and Nearctic (5%). Reptiles specimens come from 247 lo- calities of 15 states and 15 regions and 36 provinces of Italy. In the collection 46,2% of European species and 91,5% of the Italian ones are represented. The most represen- ted zoogeographic region are the Palaearctic with 89% of species in the collection, followed by the Ethiopian (14%), the Eastern (13%), Neotropical (11%) and the Nearctic (7%); australian and malagasy species do not exceed 3%. The herpetology collection was the subject of taxonomic and biogeographic analysis.
Megia-Palma, R. & Jorge, A. & Reguera, S. (2018) -
Melanic pigments play a key role in the coloration of animals, but the type of melanin pigment in black, brown, and blue colored scales of Squamata has not been studied. Based on research on birds and mammals, we may expect that pheomelanin is the majority pigment in brownish colorations and eumelanin is the majority pigment in black and blue colorations of Squamata. To characterize the pigments that underlie the melanin-based colorations of lizards, we analyzed the skin of nine genera of lacertids using dispersive Raman spectroscopy. Our results suggest that no prediction can be made on the type of pigmentary melanin present in the skin of the lacertids based alone on the hue of the sample. Indeed, brownish patterns in the skin of Psammodromus, Gallotia, Acanthodactylus, and Algyroides lizards presented both chemical forms of melanin: eumelanin and pheomelanin. Therefore, pheomelanogenesis might be an ancient characteristic within Lacertidae, because it was detected in genera in the Lacertini, Eremini, and Gallotini. Raman spectra of melanic-based patterns of genus Zootoca and ultraviolet (UV)-blue patches of Podarcis, Iberolacerta, Lacerta, and Timon lizards suggested that eumelanin is the majority pigment in these patches. Raman spectroscopy is a suitable nondestructive technique useful to identify melanin forms in the skin of lizards, and it demonstrated that pheomelanin is synthesized by Squamata.
Megia-Palma, R. & Martinez, J. & Cuervo, J.J. & Belliure, J. & Jiménez-Robles, O. & Gomes, V. & Cabido, C. & Pausas, J.G. & Fitze, P.S. & Martin, J. & Merino, S. (2018) -
Current and past parasite transmission may depend on the overlap of host distributions, potentially affecting parasite specificity and co-evolutionary processes. Nonetheless, parasite diversification may take place in sympatry when parasites are transmitted by vectors with low mobility. Here, we test the co-speciation hypothesis between lizard final hosts of the Family Lacertidae, and blood parasites of the genus Schellackia, which are potentially transmitted by haematophagous mites. The effects of current distributional overlap of host species on parasite specificity are also investigated. We sampled 27 localities on the Iberian Peninsula and three in northern Africa, and collected blood samples from 981 individual lizards of seven genera and 18 species. The overall prevalence of infection by parasites of the genus Schellackia was ∼35%. We detected 16 Schellackia haplotypes of the 18S rRNA gene, revealing that the genus Schellackia is more diverse than previously thought. Phylogenetic analyses showed that Schellackia haplotypes grouped into two main monophyletic clades, the first including those detected in host species endemic to the Mediterranean region and the second those detected in host genera Acanthodactylus, Zootoca and Takydromus. All but one of the Schellackia haplotypes exhibited a high degree of host specificity at the generic level and 78.5% of them exclusively infected single host species. Some host species within the genera Podarcis (six species) and Iberolacerta (two species) were infected by three non-specific haplotypes of Schellackia, suggesting that host switching might have positively influenced past diversification of the genus. However, the results supported the idea that current host switching is rare because there existed a significant positive correlation between the number of exclusive parasite haplotypes and the number of host species with current sympatric distribution. This result, together with significant support for host-parasite molecular co-speciation, suggests that parasites of the genus Schellackia co-evolved with their lizard hosts.
Méhely, L. (1904) -
Méhely, L. (1905) -
Mehely, L. (1907) -
Méhely, L. (1909) -
Meiri, S. (2008) -
Aim Body size is instrumental in influencing animal physiology, morphology, ecology and evolution, as well as extinction risk. I examine several hypotheses regarding the influence of body size on lizard evolution and extinction risk, assessing whether body size influences, or is influenced by, species richness, herbivory, island dwelling and extinction risk.
Methods I used literature data and measurements of museum and live specimens to estimate lizard body size distributions.
Results I obtained body size data for 99% of the world`s lizard species. The body size–frequency distribution is highly modal and right skewed and similar distributions characterize most lizard families and lizard assemblages across biogeographical realms. There is a strong negative correlation between mean body size within families and species richness. Herbivorous lizards are larger than omnivorous and carnivorous ones, and aquatic lizards are larger than non-aquatic species. Diurnal activity is associated with small body size. Insular lizards tend towards both extremes of the size spectrum. Extinction risk increases with body size of species for which risk has been assessed.
Main conclusions Small size seems to promote fast diversification of disparate body plans. The absence of mammalian predators allows insular lizards to attain larger body sizes by means of release from predation and allows them to evolve into the top predator niche. Island living also promotes a high frequency of herbivory, which is also associated with large size. Aquatic and nocturnal lizards probably evolve large size because of thermal constraints. The association between large size and high extinction risk, however, probably reflects a bias in the species in which risk has been studied.
Meiri, S. & Bauer, A.M. & Chirio, L. & Colli, G.R. & Das, I. & Doan, T.M. & Feldman, A. & Herrera, F.-C. & Novosolov, M. & Pafilis, P. Pincheira-Donoso, D. & Powney, G. & Torres-Carvajal, O. & Uetz, P. & Van Damme, R. (2013) -
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.
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
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.
Meiri, S. & Brown, J.H. & Sibly, R.M. (2011) -
Aim We provide a new quantitative analysis of lizard reproductive ecology. Comparative
studies of lizard reproduction to date have usually considered life-history
components separately. Instead, we examine the rate of production (productivity
hereafter) calculated as the total mass of offspring produced in a year. We test
whether productivity is influenced by proxies of adult mortality rates such as
insularity and fossorial habits, by measures of temperature such as environmental
and body temperatures, mode of reproduction and activity times, and by environmental
productivity and diet. We further examine whether low productivity is
linked to high extinction risk.
Methods We assembled a database containing 551 lizard species, their phylogenetic
relationships and multiple life history and ecological variables from the literature.
We use phylogenetically informed statistical models to estimate the factors
related to lizard productivity.
Results Some, but not all, predictions of metabolic and life-history theories are
supported. When analysed separately, clutch size, relative clutch mass and brood
frequency are poorly correlated with body mass, but their product – productivity –
is well correlated with mass. The allometry of productivity scales similarly to
metabolic rate, suggesting that a constant fraction of assimilated energy is allocated
to production irrespective of body size. Island species were less productive than
continental species.Mass-specific productivity was positively correlated with environmental
temperature, but not with body temperature. Viviparous lizards were
less productive than egg-laying species. Diet and primary productivity were not
associated with productivity in any model. Other effects, including lower productivity
of fossorial, nocturnal and active foraging species were confounded with
phylogeny. Productivity was not lower in species at risk of extinction.
Main conclusions Our analyses show the value of focusing on the rate of annual
biomass production (productivity), and generally supported associations between
productivity and environmental temperature, factors that affect mortality and the
number of broods a lizard can produce in a year, but not with measures of body
temperature, environmental productivity or diet.
Melendro, J. & Gisbert, J. (1976) -
Mencia, A. & Ortega, Z. & Pérez-Mellado, V. (2016) -
We conducted an experiment on chemical discrimination of two saurophagous snakes (the smooth snake, Coronella austriaca and the Seoane`s viper, Vipera seoanei) as well as the aquatic Natrix maura, by the mountain lizard Iberolacerta galani. Using terraria, 24 lizards were exposed to scents by the three snakes as well as an odourless control. We quantified fourteen behavioural variables, twelve of which significantly differed among treatments. Lizards are able to recognise the scents of both predatory snakes, and react to them with intense antipredatory responses. The antipredatory behaviour found in I. galani was similar for the scents of the two different predatory snakes, despite differences in their foraging behaviour. The behaviour displayed by lizards confronted with chemical cues suggests an adaptation to minimise the likelihood of being attacked.
Mertens, R. (1929) -
Mertens, R. (1937) -
Mertens, R. & Müller, L. (1928) -
Mertens, R. & Wermuth, H. (1960) -
Mészáros, B. & Jordán, L. & Bajer, K. & Martin, J. & Török, J. & Molnár, O. (2019) -
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.
Michelot, M. & Martinez Rica, J.P. (1989) -
Moigne, C. le & Jailloux, A. (2013) -
Monasterio, C. & Salvador, A. & Díaz, J.A. (2010) -
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.
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.
Monasterio, C. & Salvador, A. & Iraeta, P. & Díaz, J.A. (2009) -
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.
Monasterio, C. & Seixas, V.A. & Beukema, W. & Araújo, M.B. (2014) -
Monasterio, C. & Shoo, L.P. & Salvador, A. & Siliceo, I. & Diaz, J.A. (2011) -
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.
Monasterio, C. & Verdu-Ricoy, J. & Salvador, A. & Diaz, J.A. (2016) -
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.
Moreira, P. & Almeida, A. & Brito e Abreu, F. & Paulo, O. & Rosa, H. & Crespo, E.G. (2001) -
Moreira, P.L. (1996) -
Moreira, P.L. (2002) -
Moreira, P.L. & Almeida, A.P. & Rosa, H.D. & Paulo, O.S. & Crespo, E.G. (1999) -
Moreira, P.L. & Barata, M. (2005) -
Infertile and non-viable fertile eggs within a reptile clutch may decrease the incubation success of the remaining eggs, as (1) opportunistic pathogens may use the nutrient resources provided by dead eggs to colonize the clutch and spread to and kill viable eggs; and (2) odours released by spoilt eggs may attract predators to the clutch. These hypotheses were tested on the Iberian rock lizard (Lacerta monticola) by comparing the incubation success of fertile eggs between clutches composed solely of fertile eggs and clutches containing a small number of dead eggs. In a laboratory experiment, fungi (Fusarium sp. and Gliocladium sp.) colonized both infertile eggs and fertile eggs that died during incubation and thereafter spread to and killed adjacent eggs. In addition, offspring hatched earlier from fungal infected eggs than from non-infected eggs. The former were smaller and lighter than the latter, as they hatched before using the full egg yolk content. Results from a field experiment did not corroborate the fungal pathogenic effects observed in the laboratory nor did they confirm that clutches containing dead eggs suffer higher predation. Despite the inconclusive results regarding the role of fungal pathogens in nature, the present study indicates that this subject deserves further investigation in reptiles.
Moreira, P.L. & Birkhead, T.R. (2004) -
Iberian Rock Lizard (
, Boulenger 1905) males produce copulatory
plugs that harden and adhere ﬁrmly inside the female cloaca immediately after copu-
lation and occlude both oviductal openings.
To determine whether plugs reduce the chance of females being inseminated by rival
males, two hypotheses were tested: (i) that plugs reduce female attractiveness and/or
receptivity to rival males; and (ii) that plugs function as ‘chastity belts’.
There was no evidence to support either hypothesis. Neither male initiation of
copulations and female termination of male copulation attempts, nor male insemina-
tion success differed signiﬁcantly between trials in which males were presented with
females without a plug and with females with a plug deposited by a different male 1, 3
and 6 or more hours earlier, nor between females without and with a plug.
Several hypotheses for the function of Iberian Rock Lizard plugs are discussed.
Since rival males were frequently able to displace a plug in the female cloaca, our results
are consistent with a ‘sperm protection’ function of male postcopulatory mate guarding
in squamate reptiles, but this remains to be tested.
Male Iberian rock lizards (Lacerta monticola)
produce copulatory plugs that adhere firmly inside the
female cloaca and occlude both oviducts. These plugs do
not prevent rival male insemination, as they neither reduce
female attractiveness or receptivity to rival males
nor do they function as “chastity belts”. Prior to copulation,
males bite various regions of the female body, including
the cloacal region. A previous plug in the female
cloaca is expelled only after rival male intromission.
We hypothesized that: (1) such male pre-copulatory behaviours
function to loosen plug adherence to the female
cloaca, thereby facilitating intromission, and that (2) the
hemipenis plays a role in displacing a previous plug prior
to the delivery of a new plug. Neither of these hypotheses
was supported. Instead, our results indicate that rival
males can intromit the hemipenis past a previous plug in
the female cloaca and deliver their own plug underneath
it. Consequently, previous plugs are pushed away from
the oviductal openings and even dislodged from the female
cloaca. Copulation duration was determined both by
the time used by males to deliver a plug and by the fact
that males prolonged copulation beyond plug delivery.
There seemed to be sexual conflict over prolonged copulation,
which was resolved by the male/female headlength
ratio. The adaptive value of Iberian rock lizard
copulatory plugs and prolonged copulation in Iberian rock
lizards is discussed in the context of sperm competition.
Moreira, P.L. & Lopez, P. & Martin, J. (2006) -
Socially dominant males often signal their status to rival males and/or females. We tested the hypotheses that Lacerta monticola femoral gland secretions and copulatory plugs convey chemical information about male identity and dominance status. We estimated male dominance status by staging male-male agonistic encounters in a neutral arena. We then conducted two experiments to compare male tongue-flick behavior toward chemical stimuli consisting of cotton swabs bearing (1) deionized water (control), the lizard`s own femoral secretions, and the femoral secretions of another male and (2) phosphate-buffered saline solution (control), the lizard`s own plug products, and the plug products of another male. Results indicate that males discriminated their own femoral secretions and plugs from those of other males. They also discriminated morphological attributes of other males that were associated with dominance status based on chemical cues arising from femoral secretions and discriminated the dominance status of other males based on chemical cues arising from the plugs. Femoral secretions that convey information about male identity and dominance status may be hypothesized to function in the establishment of L. monticola dominance hierarchies through scent-marking of territories. We suggest that copulatory plugs and femoral secretions may allow males to scent-mark the female body and postulate that this behavior may influence male and female reproductive decisions under selective pressures of sperm competition.
Moreira, P.L. & López, P. & Martin, J. (2008) -
Faeces play a role in intraspecific chemical communication in many vertebrates, including lizards. Here, we hypothesised that juvenile Iberian rock lizards (Lacerta (=Iberolacerta) monticola) use substrate-borne conspecific faecal chemicals to assess the social environment and base their spatial decisions. To test this hypothesis, we prepared chemical stimuli by dissolving faeces in an organic solvent (dichloromethane, DCM) and conducted chemosensory trials where we quantified latency to the first tongue flick (TF) and TF rates when young captive-borne juveniles were placed in an arena compartment whose substrate was labelled by (1) a control (DCM), (2) their own faecal chemicals, (3) faecal chemicals of another juvenile, (4) faecal chemicals of an adult female or (5) faecal chemicals of an adult male. Following TF observations, we removed a partition that separated the labelled compartment from a chemically unlabelled one, and recorded for 5 min when juveniles first crossed to the unlabelled compartment and total time spent in the labelled compartment. Each juvenile was tested with all stimuli in a randomised order. In addition, juveniles belonged to different families and were unrelated to and unfamiliar with faeces donors. Taken together, TF and spatial responses toward stimuli indicate that juveniles discriminated between faecal chemicals of conspecific juveniles, adult females and males, and that they avoided remaining in substrates labelled by adult male faecal chemicals. We suggest that juveniles assess the social environment based on conspecific faecal chemicals thus avoiding aggression and cannibalistic risks undertaken from encounters with adult males.
Moreira, P.L. & Nunes, V.L. & Martín, J. & Paulo, O.S. (2007) -
Sperm competition selects for opposing male
defensive and offensive reproductive traits, and its outcome
may be determined by the effectiveness to which one trait
has evolved to out-compete the other. We tested the
effectiveness of a first male plug physical interference with
a second male insemination (defence) vs the effectiveness
of plug and associated sperm displacement by a second
male (offence) on the outcome of sperm competition in
Iberian rock lizards. We conducted a double mating experiment
where we compared the proportion of eggs per
clutch fertilised by the same second males (against the same
first males) when they copulated with females 30 min (first
male plug adhered firmly inside the female cloaca) and 4 h
(first male plug loosely adhered or shed from the female
cloaca) after first males. We found that second males
fertilised the majority of the eggs per clutch in the 30-min
treatment, whereas fertilisations were equally shared between
the two males in the 4-h treatment. These results
show that plugs have little defensive effectiveness, and
thus, do not assure high first male fertilisation success.
Instead, sperm displacement appears to be associated with
plug displacement. That is, because sperm embedded in
first male plugs, and displaced from competition for fertilisations by second males, is expected to increase in
number with decreasing time allowed for female sperm
transport, second males thus enjoy higher fertilisation
success. This study shows that offensive plug displacement
out-competes plug defensive role in Iberian rock lizards.
Moreover, it reveals sperm displacement as a novel sperm
competition mechanism in reptiles.
Moreira, P.L. & Paulo, O.S. (2008) -
Mouret, V. & Guillaumet, A. & Cheylan, M. & Pottier, G. & Ferchaud, A.-L. & Crochet, P.-A. (2011) -
Aim The genetic impact of Quaternary climatic fluctuations on mountain endemic species has rarely been investigated. The Pyrenean rock lizard (Iberolacerta bonnali) is restricted to alpine habitats in the Pyrenees where it exhibits a highly fragmented distribution between massifs and between habitats within massifs. Using mitochondrial DNA markers, we set out: (1) to test whether several evolutionary units exist within the species; (2) to understand how the species persisted through the Last Glacial Maximum and whether the current range fragmentation originates from distribution shifts after the Last Glacial Maximum or from more ancient events; and (3) to investigate whether current mitochondrial diversity reflects past population history or current habitat fragmentation.
Location The Pyrenees in south-western France and northern Spain.
Methods We used variation in the hypervariable left domain of the mitochondrial control region of 146 lizards collected in 15 localities, supplemented by cytochrome b sequences downloaded from GenBank to cover most of the species’ distribution range. Measures of population genetic diversity were contrasted with population isolation inferred from topography. Classical (F-statistics) and coalescence-based methods were used to assess the level of gene flow and estimate divergence time between populations. We used coalescence-based simulations to test the congruence of our genetic data with a scenario of simultaneous divergence of current populations.
Results Coalescence-based analyses suggested that these peripheral populations diverged simultaneously at the end of the last glacial episode when their habitats became isolated on mountain summits. High mitochondrial diversity was found in peripheral, isolated populations, while the populations from the core of the species’ range were genetically impoverished. Where mitochondrial diversity has been retained, populations within the same massif exhibited high levels of genetic differentiation.
Main conclusions As suggested for many other mountain species, the Pyrenean rock lizard survived glacial maxima through short-distance range shifts instead of migration or contraction in distant southern refugia. Most of the main Pyrenean range has apparently been re-colonized from a single or a few source populations, resulting in a loss of genetic diversity in re-colonized areas. As a result, current levels of intra-population mitochondrial diversity are better explained by post-glacial population history than by current habitat fragmentation. Genetic population differentiation within massifs implies severe reduction in female-mediated gene flow between patches of habitats.
Mršić, N. (1978) -
Mudde, P. (1991) -
A BLACK SPECIMEN
OF LACERTA MONT/COLA
Isolation and environmental factors can result in special colored populations of lizards on moun- taintops and islands. Near Lago Ercina in the Pi- cos d`Europa a single all black specimen ofLacer- ta monticola was found. Although this specimen could be infertile, ill or not capable of getting a suitable territorium, the author nevertheless sug- gests the possibility of the development of a black population in about ten years time.
Müller, L. & Hellmich, W. (1937) -
Muratet, J. (2015) -
Nature Midi Pyreneés (2010) -
Naulleau, G. (1980) -
Naulleau, G. (1990) -
Nikolsky, A.M. (1915) -
Nores Quesada, C. & Garcia-Rovés González, P. (2007) -
Odierna, G. & Aprea, G. & Arribas, O. & Capriglione, T. & Caputo, V. (1996) -
The karyology of two taxa represented in the mountain herpetofauna: Discoglossus Otth, 1837 and Iberian Archaeolacerta Méhely, 1909 -- Discoglossus and Iberian Archaeolacerta represent good models in studying the roles played by chromosomal variations during evolution. In Discoglossus chromosomal rearrangements appear to have essentially involved the localisation and/or composition of heterochromatin and, at a lower degreee, the chromosome morphology. Kariologic data on the Iberian Archaeolacerta showed that L. monticola is a polytypical taxon including L. monticola sensu stricto and the L. bonnali complex. In the former taxon there are 2n=36 all acrocentric chromosomes, and in the latter there are 2n=24-26 chromosomes, 10-12 of which biarmed.
Odierna, G. & Aprea, G. & Arribas, O. & Capriglione, T. & Olmo, E. (2001) -
Odierna, G. Aprea, G. Arribas, O. Capriglione, T. & Olmo, E. (1996) -
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.
Oefinger, B. & Oefinger, P. (2010) -
Oefinger, B. & Oefinger, P. (2011) -
Oefinger, B. & Oefinger, P. (2013) -
Oefinger, B. & Oefinger, P. (2014) -
Oefinger, B. & Oefinger, P. (2016) -
Oefinger, B. & Oefinger, P. (2019) -
Olmo, E. & Aprea, G. & Arriba, O.J. & Barucca, M. & Canapa, A. & Capriglione, T. & Bosch, H. in den & Odierna, G. (2004) -
Olmo, E. & Odierna, G. & Capriglione, T. (1987) -
The occurrence and form of sex chromosomes were investigated with the aid of C-banding and 4′-6-diamidino-2-phenylindole (DAPI) staining in 13 species of lacertid lizards. The results obtained show the presence in five species of a female heterogamety in which the two sex chromosomes have the same shape and size, but the W differs from the Z in being almost entirely heterochromatic. This condition is clearly similar to that found in some snakes and considered to be an early stage of differentiation of sex chromosomes by Singh et al. (1976, 1980). A more evolved condition may be that found in three other species in which the W is distinctly smaller than the Z. A third situation is that found in all Podarcis species which, even though they are considered to be among the more evolved species in the family, possess two sex chromosomes that are indistinguishable. In general, the situation in lacertids may be compatible with the hypothesis of sex chromosome evolution put forward by Singh et al. (1976, 1980). However a differentiation mechanism of this kind does not seem to be well established in lacertids, and is probably not the only mechanism that is in operation in this family.
Ortega Diago, Z. (2016) -
All aspects of the biology of ectotherms are influenced by their body temperatures, which, in turn, depend heavily on the temperature of the habitat in which they live. High mountain habitats impose severe constrains, being the short length of the daily and the annual activity periods, together with the low ambient temperatures, the most relevant to the thermal biology of animals. In this thesis, we address the thermal biology of five species of the high mountain lizard genus Iberolacerta: I. cyreni, I. galani, I. aurelioi, I. bonnali, and I. monticola. We have studied the thermal sensitivity of all of them, and the degree to which they are able to regulate their body temperatures behaviorally, assessing accuracy of thermoregulation, thermal quality of the habitat, and effectiveness of thermoregulation. To do this, we have measured body temperatures of activity, operative temperatures as null hypothesis of thermoregulation, and temperatures selected in laboratory thermal gradient. Additionally, in particular cases of study, we have evaluated: (1) the role of behavioral thermoregulation to buffer the impact of climate change in body temperatures, (2) the role of the seasonal shift of the preferred temperatures range in the effectiveness of thermoregulation, (3) the partition of thermal resources of two syntopic species, and (4) the impact of wind speed in the effectiveness of thermoregulation. General results show that Iberolacerta species are cold-specialists, as well as excellent thermoregulators. In the case of I. cyreni, we have shown that lizards are buffering the impact of climate change by means of behavioral thermoregulation. With regard to I. galani, we have found that the shift of its preferred temperatures range between spring and summer entails a more effective thermoregulation for each season. In addition, we have studied the partition of the thermal niche between I. galani and Podarcis bocagei in sympatry. Finally, we have shown that an increase in wind speed significantly affects the effectiveness in thermoregulation I. aurelioi. All results of the thesis have an added importance due to the current climate change scenario, which is particularly threatening these species of lacertids of the Iberian mountains. Therefore, we have discussed all results in the light of this unfortunate phenomenon, with the hope that contributions to the knowledge of the thermal biology of ectotherms will be useful to prevent their extinction in the medium term.
Ortega, Z. & Mencía, A. & Pérez-Mellado, V. (2016) -
We studied, at 2200 m altitude, the thermal biology of the Pyrenean rock lizard, Iberolacerta bonnali, in glacial cirque of Cotatuero (National Park of Ordesa, Huesca, Spain). The preferred thermal range (PTR) of I. bonnali indicates that it is a cold-adapted ectotherm with a narrow PTR (29.20-32.77 °C). However, its PTR (3.57 °C) is twice as wide as other Iberolacerta lizards, which may be explained by its broader historical distribution. The studied area is formed by a mosaic of microhabitats which offer different operative temperatures, so that lizards have, throughout their entire daily period of activity, the opportunity to choose the most thermally suitable substrates. I. bonnali achieves an effectiveness of thermoregulation of 0.95, which makes it the highest value found to date among Lacertidae, and one of the highest among lizards. Their relatively wide distribution, their wider PTR, and the excellent ability of thermoregulation, would make I. bonnali lizards less vulnerable to climate change than other species of Iberolacerta. Thanks to its difficult access, the studied area is not visited by a large number of tourists, as are other areas of the National Park. Thus, it is a key area for the conservation of the Pyrenean rock lizard. By shuttling between suitable microhabitats, lizards achieve suitable body temperatures during all day. However, such thermally suitable microhabitats should vary in other traits than thermal quality, such as prey availability or predation risk. Hence, it seems that these not-thermal traits are not constraining habitat selection and thermoregulation in this population. Therefore, future research in this population may study the causes that would lead lizards to prioritize thermoregulation to such extent in this population.
Mountain lizards are highly vulnerable to climate change, and the continuous warming
of their habitats could be seriously threatening their survival. We aim to compare the
thermal ecology and microhabitat selection of a mountain lizard, Iberolacerta galani,
and a widely distributed lizard, Podarcis bocagei, in a montane area. Both species
are currently in close syntopy in the study area, at 1,400 m above the sea level. We
determined the precision, accuracy and effectiveness of thermoregulation, and the
thermal quality of habitat for both species. We also compared the selection of thermal
microhabitats between both species. Results show that I. galani is a cold-adapted
thermal specialist with a preferred temperature range of 27.929.7 C, while P. bocagei
would be a thermal generalist, with a broader and higher preferred temperature range
(30.134.5 C). In addition, I. galani selects rocky substrates while P. bocagei selects
warmer soil and leaf litter substrates. The thermal quality of the habitat is higher for P.
bocagei than for I. galani. Finally, P. bocagei achieves a significantly higher effectiveness
of thermoregulation (0.87) than I. galani (0.80). Therefore, these mountain habitat
conditions seem currently more suitable for performance of thermophilic generalist
lizards than for cold-specialist lizards.
Ortega, Z. & Mencia, A. & Pérez-Mellado, V. (2016) -
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.
Thermal biology of lizards affects their overall physiological performance. Thus, it is crucial to study how abiotic constraints influence thermoregulation. We studied the effect of wind speed on thermoregulation in an endangered mountain lizard (Iberolacerta aurelioi). We compared two populations of lizards: one living in a sheltered rocky area and the other living in a mountain ridge, exposed to strong winds. The preferred temperature range of I. aurelioi, which reflects thermal physiology, was similar in both areas, and it was typical of a cold specialist. Although the thermal physiology of lizards and the structure of the habitat were similar, the higher wind speed in the exposed population was correlated with a significant decrease in the effectiveness thermoregulation, dropping from 0.83 to 0.74. Our results suggest that wind reduces body temperatures in two ways: via direct convective cooling of the animal and via convective cooling of the substrate, which causes conductive cooling of the animal. The detrimental effect of wind on thermoregulatory effectiveness is surprising, since lizards are expected to thermoregulate more effectively in more challenging habitats. However, wind speed would affect the costs and benefits of thermoregulation in more complex ways than just the cooling of animals and their habitats. For example, it may reduce the daily activity, increase desiccation, or complicate the hunting of prey. Finally, our results imply that wind should also be considered when developing conservation strategies for threatened ectotherms.
The León rock lizard, Iberolacerta galani, lives in isolated mountains of Spain.We studied the seasonal changes in the thermal biology of I. galani between spring and summer. We calculated precision, accuracy andeffectiveness of thermoregulation and the habitat thermal quality for spring, and compared with the values of summer. In addition, we studied how the shift in the thermal preferences of lizards would contribute to achieve a higher effectiveness of thermoregulation. Thermal preferences of León rock lizards are among the lowest in lacertids, and are also very narrow, maintaining the narrowness among seasons. As for summer (27.90–29.70 °C, mean value =28.76 °C), the thermal preferences of I. galani are also low in spring (29.60–31.10 °C, mean value =30.38 °C), supporting the idea that this species is adapted to cold environments. The habitat thermal quality is lower in spring (10.99 °C) than in summer (9.36 °C), while the effectiveness of thermoregulation is higher in spring (0.92) than in summer (0.80). We found that the seasonal shift in thermal preferences contributes significantly to enhance the effectiveness of thermoregulation in both seasons, more in spring (0.45 °C) than in summer (0.16 °C). Because I. galani inhabits isolated mountains, where the activity period is reduced from April to October, we hypothesize that the observed adaptation of the thermal preferences, which enhance thermoregulation to a larger extent in spring, may evolved to maximize performance during the reproductive season.
Ortega, Z. & Mencia, A. & Pérez-Mellado, V. (2017) -
We studied the thermal ecology of the montane Iberian rock lizard, Iberolacerta monticola, in the western area of its distribution at the Serra da Estrela (Portugal). We calculated the precision of thermoregulation and the indices of thermal quality of the habitat, and accuracy and effectiveness of thermoregulation. To complete the study of the thermal ecology, we assessed the relationships between body and environmental temperatures, and we described the thermal and spatial heterogeneity of the habitat. Our results indicate that the Iberian rock lizard is a cold-specialist, with a preferred temperature range between 29.80 and 31.60 °C. Thus, precision of thermoregulation is 1.8 °C, which is a normal range in thermal specialists, like other species of the genus Iberolacerta. This result is important because being thermal specialists and living in mountaintops make Iberian mountain lizards particularly vulnerable to global warming. The habitat of I. monticola at the Serra da Estrela is formed of microhabitats offering different operative temperatures, which allows lizards to select the most suitable for thermoregulation at any time of the day. Iberian rock lizards achieve an effectiveness of thermoregulation of 0.86, thanks to careful thermoregulatory behaviour. Rocky microhabitats occupy more than 50% of its habitat, so is probable that lizards are selecting rocks to warm themselves faster, minimising the costs of thermoregulation. A possible thigmothermic component of this kind would be unique among the species of Iberolacerta.
Ortega, Z. & Mencia, A. & Pérez-Mellado, V. (2018) -
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.
Osojnik, N. (2012) -
Osojnik, N. & Zagar, A. & Carretero, M.A. & Garcia-Munoz, E. & Vrezec, A. (2013) -
The lizards Podarcis muralis and Iberolacerta horvathi display an overall similarity in morphology and ecology. Although they are widely sympatric in part of their distribution range of Slovenia, areas of altitudinal segregation have been observed. Ecophysiological traits such as preferred body temperatures (Tp) and rates of evaporative water loss (EWL) are some of the factors likely to be subjacent to this pattern. We expect the species that is more abundant in higher altitudes to have lower Tp and higher EWL (due to lower environmental temperatures at higher elevations) than the species that is more abundant in lowlands. We conducted lab experiments to determine whether intra- and interspecific variation in these two physiological traits exists. Contrary to expectations, the means of Tp were similar between the species, but their seasonal dynamics differed, which indicates that thermal accuracy might be more important. Species differed in seasonal (and daily) variation of Tp, rising from spring to summer more in P. muralis than in I. horvathi. Comparing instantaneous EWL, our results showed interspecific differences with higher values in P. muralis than in I. horvathi. Throughout a 12-h period, the accumulative EWL was also always higher for P. muralis than for I. horvathi, with the greatest differences occurring after 9–12 h of the experiment. Further studies should focus on investigating if these divergent physiological characteristics of both species are associated with their habitat use (in terms of thermal and water environment) and species interaction.
Osojnik, N. & Zagar, A. & Vrezec, A. & Carretero, M.A. (2010) -
Distribution of the genera Podarcis and lberolacerta in Europe have been hypothesized to derive from current and past interactions. Among the putative factors involved, thermal ecology is expected to play a determinant role. As a first step in this research, here we perform a comparative analysis of the preferred temperatures (Tp) by representatives of both genera living in sympatry. Because Tp carries substantial phylogenetic inertia, we expect derive predictions for other similar species tandems. Podarcis muralis and lberolacetfa horvathi display overall similarity in morfometry, coloration and ecology. While P. muralis is a widespread species in Europe occupying a variety of microhabitats, including urban areas, I. horvathi is endemic to Southern Alps and Dinaric Mountains living in rocky habitats but never near human settlements. Sympatry of both species has been recorded in Italy, Austria and, recently, Slovenia. Slovenian I. horvathi is more found at higher altitudes whereas I? muralis shows the opposite trend suggesting higher Tp; To test our hypothesis we have preformed experiments in laboratory thermo gradients (20 -50°C; measurements at 11 hourly intervals) with specimen form a sympatric area from the KoEevska region (SE Slovenia). Preliminary results with males indicate that, contrary to the expectations, Tp were slightly higher in I. horvathi than in P. muralis. However, such differences were only significant in the central hours of the day, hence, P. muralis being more variable daily. This pattern, suggesting more importance of thermal tolerance than thermoregulatory set point, is to be tested by further field studies.
Palacios, F. & Salvador, A. (1974) -
Palanca, Antonio & Rey, Jorge & Riobó, Antonio & Vences, Miguel (1997) -
Palaus, J. (1974) -
Parent, G.H. (1981) -
ventaire exhaustif des espèces indigènes et naturalisées, présentes e n France et au Benelux : 85 taxons, relevant de 72 espèces, sont cités, mais deux autre s taxons, dont le statut reste à définir, sont mentionnés au n os 31 (Rana esculenta) et a u n° 53 (Podarcis muralis) et huit autres au n` 56 (Podarcis muralis tiliguerta) de l`inven- taire. On signale les zones à prospecter par priorité et les problèmes taxonomiques à résoudre d`urgence .
Les aires sont définies par rapport aux provinces ou aux départements : des cartes d`aire provisoires, en France, sont données pour 33 taxons.
En ce qui concerne la liste des taxons relatifs à la France, les modifications suivante s sont à noter :
1°) taxons admis : Triturus alpestris apuanus, Acanthodactylus erythrurus cf. erythru- rus, Lacerta agilis garzoni, Podarcis hispanica hispanica, Podarcis muralis merremia ,
Vipera latastei latastei, Vipera seoanei ;
2.) deux intergrades de Salamandra salamandra sont retenus :
30) taxons refusés : Rana iberica, Coronella austriaca fitzingeri, Natrix tessellata
4.) tombent en synonymie les deux taxons suivants : Podarcis muralis occidentalis
(= P. m. muralis) et P. m. calbia (= P. m. oyensis).
En outre, liste des espèces introduites, liste des espèces citées par erreur et list e
des taxons à rechercher . Quelques données inédites concernent le nord de l`Espagne .
Pavlicev, M. & Mayer, W. (2009) -
Lacertinae is one of the three lacertid lizard subfamilies with a geographical distribution confined to the
Palaearctic. Several past attempts to reconstruct its phylogeny resulted in unresolved bush-like topologies.
We address the question of whether the lack of resolution is due to insufficient data or whether this
lack reflects a rapid succession of speciation events. We analyzed four partial and one complete gene
sequences from mitochondrial and nuclear genomes, totalling roughly 3600 bp. We included 29 species
representing all 19 genera suggested in recent revision of Lacertinae [Arnold, E.N., Arribas, O., Carranza, S.,
2007. Systematics of the palaearctic and oriental lizard tribe Lacertini (Squamata: Lacertidae: Lacertinae),
with descriptions of eight new genera. Zootaxa 1430, 1–86]. The resulting phylogeny, first, corroborates
monophyly at the genus level for the suggested genera, as well as the finding that Atlantolacerta andreanskyi,
until recently part of Lacertinae, belongs to the subfamily Eremiadinae. Second, we find that
increasing the sequence length and combining multiple nuclear and mitochondrial sequences did not
resolve the polytomy, suggesting that the inferred topology indicates a multiple cladogenesis within a
short geological period, rather than a methodical artefact. Divergence time estimates, based on previous
estimates of several node ages, range from 13.9 to 14.9 million years for the radiation event, however
with very broad confidence interval. To associate the radiation with a narrower geological time we consider
palaeogeographic and palaeoclimatic data, assuming that the Lacertinae probably evolved in Central
Europe andWAsia after the collision of Africa and Eurasia. We suggest that this radiation may date to the
late Langhian (ca. 14–13.5 million years) when geological events caused abrupt changes in regional
water–land distribution and climate, offering a window of distinct conditions.
Pedro Silva, J. & Toland, J. & Jones, W. & Eldridge, J. & Hudson, J. & O`Hara, E. (2009) -
Peek, R. (2015) -
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.
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.
Pérez i de Lanuza, G. & Font, E. (2016) -
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.
Pérez-Mellado, V. (1982) -
A study on taxocenosis of Lacertidae with 8 species: Lacerta lepida, Lacerta schreiberi, Lacerta montícola, Acanthodactylus erythrurus, Psammodromus algirus, Psammodromus hispanicus, Podareis hispanica and Podareis bocagei in the western part of the Sistema Central (Iberian Península) has been carried out. Most of these behave as feeding generalists but A. erythrurus seems to specialize in mymecophagy whereas L. montícola turns out to be a «forced» sthenophage. The overlap valúes in each niche dimensión suggest that the spatial dimensión and habitat preference are the most important factors. However, segregation is produced jointly due to the interaction of the three dimensions considered which act interdependently.
Pérez-Mellado, V. (1997) -
Pérez-Mellado, V. (1998) -
Pérez-Mellado, V. (2002) -
Pérez-Mellado, V. & Barbadillo, L.J. & Barahona, F. & Brown, R.P. & Corti, C. & Guerrero, F. & Lanza, B. (1993) -
Pérez-Mellado, V. & Bauwens, D. & Gil, M. & Guerrero, F. & Lizana, M. & Ciudad,M.J. (1991) -
We studied diet composition and prey selection in the lizard Lacerta monticola throughout its activity season. The most important prey groups in terms of numbers were Coleoptera, Diptera, Formicidae, and Araneae. Comparison of diet composition in the adult lizards with estimates of prey availability indicated that lizards exhibited precisely defined patterns of prey selection. Electivities were negative for prey < 3 mm in length and generally positive for larger prey, despite the higher relative abundance of the former group in the environment. Electivity scores for individual prey taxa were positively correlated with their length, and lizards preferentially consumed larger individuals belonging to the taxa containing smaller animals. Relative availability of the four major prey taxa fluctuated considerably throughout the year. In spite of these changes, monthly rates of consumption of Araneae and Diptera remained almost constant. Variations in monthly electivity scores were negatively correlated with relative availability of three (Coleoptera, Diptera, and Araneae) of the four major prey taxa. These results fit the predictions of prey-selection models based on nutrient constraints.
Pérez-Mellado, V. & Cheylan, M. & Martinez-Solano, I. (2009) -
Pérez-Mellado, V. & Corti, C. & Lo Cascio, P. (1997) -
Tail autotomy is one of the main anti-predator mechanisms of lacertid lizards, but it has been predicted that it is only retained in its full capacity when its benefits exceed its costs (Arnold, 1988). To test this hypothesis, ease of tail shedding was examined in a number of continental and insular lacertid lizard populations, each of which showed a different shedding capacity. Tails are shed more easily in those continental and insular populations where there is a greater probability of predation. In insular populations not subjected to strong predation, the tail tends to be retained. The relationship of these findings to insular Mediterranean lizard populations and to the extinction of the Balearic lizard, Podarcis lilfordi are discussed.
Pérez-Mellado, V. & Gil, M.J. & Guerrero, F. & Pollo, C. & Rodriguez, E. & Marco,A. & Liana, M. (1988) -
Pérez-Mellado, V. & Marquez, R. & Martinez-Solano, I. (2009) -
Pérez-Mellado, V. & Sa-Sousa, P. & Marquez, R. & Martinez-Solano, I. (2009) -
Polo, V. & López, P. & Martin, J. (2005) -
When prey take refuge to reduce predation risk, they forfeit time for other activities. They may also pay a physiological cost. In particular, optimal regulation of body temperature is essential for ectotherms. Qualitative models predict that lizards have to balance anti-predatory decisions in relation to thermal conditions of the refuge and predation risk when deciding when to resume activity. However, these models are not general and empirical tests of their assumptions are scarce. We modified previous models to include the case of a high and maintained level of predation risk. The predictions of the model were further investigated in a laboratory experiment using male Iberian rock lizards. The same level of predation risk was displayed in two treatments in which temperature inside the refuge was high or low, and in the mating or the post-reproductive season. As predicted, lizards increased successive emergence times - an increase that was not linear but accelerating - and they had shorter emergence times when thermal costs of refuge use were higher. Nevertheless, body size of lizards and the season of the experiment had no effect on these decision rules. Our results are in line with an economical balance between costs and benefits in the decision rules controlling active versus inactive periods
Polo, V. & López, P. & Martín, J. (2011) -
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.
Pottier, G. (2007) -
Découvert en 1922 dans les Hautes-Pyrénées, le Lézard pyrénéen de Bonnal (Iberolacerta bonnali) est une espèce endémique de la ceinture alpine des Pyrénées centrales, inscrite à l’annexe 2 de la directive européenne Habitats – Faune –Flore. En France, son aire de répartition est presque entièrement située sur le territoire du Parc National des Pyrénées, et en grande partie intégrée aux sites Natura 2000 qui s’y trouvent. Souhaitant optimiserla conservation de cette espèce à forte valeur patrimoniale envers laquelle sa responsabilité est très élevée, le P.N.P. a commandé une étude visant à mieux connaître le Lézard pyrénéen de Bonnal, et comprenant les volets suivants: répartition spatiale et altitudinale, sélection d’habitat, structuration génétique des populations à différentes échelles spatiales, dynamique des populations, biométrie et relation avec les lacertidés sympatriques et syntopiques, notamment le Lézard des murailles Podarcis muralis. Après avoir présenté les caractéristiques du Lézard pyrénéen de Bonnal et du Parc National des Pyrénées, nous livrons les résultats de cette étude: ayant découvert de nombreuses localités nouvelles de l’espèce distribuées sur l’ensemble des massifs du P.N.P., nous avons pu établir que l’espèce occupe la quasi-totalité de son aire de répartition potentielle (étage alpin) sur la zone d’étude. Cependant, la structuration génétique des populations révèle un faible taux (voire une absence) de connexion, y compris entre localités proches, ce fait étant à mettre en relation avec la distribution de l’habitat favorable et la faible mobilité de l’espèce. Enfin, le Lézard des murailles apparaît comme un compétiteur. Ainsi, bien qu’occupant des zones de moyenne et haute montagne relativement reculées où l’impact des nuisances écologiques d’origine humaine apparaît a priori négligeable, le Lézard pyrénéen de Bonnal se présente comme une espèce vulnérable du fait de la fragmentation de ses populations et de la possibilité d’une compétition accrue avec le Lézard des murailles en raison du réchauffement climatique.
Pottier, G. (2008) -
Pottier, G. (2012) -
Pottier, G. (2016) -
Les Reptiles des Pyrénées traite en détail les 32 espèces protégées présentes sur l’ensemble de la chaîne (France, Espagne et Andorre). Cette faune herpétologique est le résultat d’une entreprise naturaliste combinant enquête bibliographique et reportage photographique. Elle propose une vaste synthèse de données relatives à plusieurs champs disciplinaires – systématique, taxinomie, biogéographie, écologie, biologie… – jusque-là dispersées dans plusieurs centaines d’articles et ouvrages. Les variations phénotypiques et les particularités écologiques des serpents, lézards et tortues occupant l’espace pyrénéen y sont illustrées par de très nombreuses photographies exclusivement réalisées in situ, de 2 m à 3 143 m d’altitude. Des cartes de répartition précises (mailles UTM 10 km × 10 km), basées sur des sources scientifiques, complètent et éclairent le propos biogéographique. Les espèces et sous-espèces endémiques ou subendémiques de la chaîne, de même que celles qui y ont un statut particulier (très localisées, vulnérables…), ont fait l’objet d’une attention particulière et les menaces qui pèsent sur les reptiles des Pyrénées sont largement exposées.
Cet ouvrage sera précieux pour les pyrénéistes, naturalistes et gestionnaires d’espaces naturels de la chaîne (parc national, réserves, sites Natura 2000, parcs naturels régionaux…), qui disposeront là d’une mine d’informations sur le sujet.
Pottier, G. (2018) -
Pottier, G. & Arthur, C.-P. & Weber, L. & Cheylan, M. (2013) -
Les trois lézards endémiques des Pyrénées: Iberolacerta aranica, Iberolacerta aurelioi et Iberolacerta bonnali, ont fait l`objet de plusieurs travaux de terrain sur le versant français de la chaîne depuis 1999. Ces travaux ont permis d`acquérir d`importantes données chorologiques sur ces trois espèces, en particuliers de nombreuses localités nouvelles qui modifient parfois sensiblement le patron de leur aire de répartition connue (tant horizontalement que verticalement). Ces données, en majorité inédites, sont ici compilées et commentées. Elles démontrent que, contrairement à ce qui était précédemment supposé, une part importante de l`effectif mondial de ces trois espèces (plus de la moitié dans le cas d`I. aranica) se situe sur le territoire français. La responsabilité de la France vis-à-vis de ces trois espèces s`en trouve sensiblement accrue. Le présent article, dernier d`une série de trois, est consacré au Lézard de Bonnal, Iberolacerta bonnali.
Pottier, G. & Delmas, C. & Duquesne, A. & Garric, J. & Paumier, J.-M. & Sfreddo, G. & Tessier, M. & Verge, J. (2010) -
Several field studies have focussed on the three Pyrenean endemic lizards (Iberolacerta aranica, Iberolacerta aurelioi and Iberolacerta bonnali) on the French side of the Pyrenees mountain range from 1999 to 2009. The studies allowed acquisition of important distributional data, all three species having been discovered in many new localities, sometimes considerably increasing their known distribution range (spatial and altitudinal). These data, mostly unpublished, have been compiled and are presented here. This improved knowledge demonstrates that, contrary to prior belief, a large part of the worldwide populations of these three species (a major part in the case of I. aranica) are actually located in France. This noticeably increases the conservation duty of France for these three species. The current article, first of a series of three, is devoted to Aran’s Rock Lizard, Iberolacerta aranica.
Pottier, G. & Garric, J. (2006) -
Trois localités du Lézard pyrénéen du Val d’Aran I b e ro l a c e rta aranica sont signalées en France dans le massif du Mont Valier (département de l’Ariège), situées au-delà de la limite orientale connue de l’aire de répartition de l’espèce (zone du port d’Orle et du pic de Barlonguère). La présence avérée d’I. aranica dans ce massif augmente sensiblement la surface de l’aire de répartition connue de l’espèce, et accroît le degré de responsabilité conservatoire de la France vis-à-vis de ce taxon très localisé, endémique de l’étage alpin des Pyrénées centrales.
Pozzi, A. (1966) -
Prieto Espineira, X. & Cabana, M. (2020) -
Radovanović, M. (1941) -
Radovanović, M. (1958) -
Radovanovic, M. (1964) -
Rassati, G. (2009) -
Contribution to the knowledge of the distribution of Horvath’s rock lizard Iberolacerta horvathi and Common wall lizard Podarcis muralis in the Friuli Venezia Giulia and in the Veneto regions (North-eastern Italy) – Data collected between 2002 and 2008, during an investigation aimed to fi nd unknown populations of Iberolacerta horvathi and at extending the knowledge about the distribution of Podarcis muralis in Northern Friuli Venezia Giulia and Veneto, are given. Iberolacerta horvathi has been found in 22 new localities, increasing of about 45% of those previously known in Italy, between 520 and 1825 metres a.s.l.. Podarcis muralis has been found in 103 localities, most of them in UTM squares (10X10 Km) not covered by the national atlas. In Friuli Venezia Giulia the species has been found regularly over 900 metres a.s.l. where suitable habitats are present, and locally it reaches 1800 metres a.s.l.. The two species have been found in syntopy in eight localities between 520 and 1800 metres a.s.l.. The Mount Dimon represents the highest locality where in Friuli Venezia Giulia Podarcis muralis has been found and the two lizards in Italy are in syntopy. Finally it has to be pointed out that the works for construction, maintenance and extension of walls, roads etcetera might drastically reduce the suitable habitat of the two species.
Rassati, G. (2019) -
In the last 20 years or so, Horvath’s Rock Lizard Iberolacerta horvathi has been found at over 40 new sites, allowing us to describe a more realistic distribution pattern of the species and to provide novel elements for its understanding. In Italy, the species is distributed only in the north-eastern extremity in 37 UTM squares. Records in the Carnic Prealps and in Veneto have greatly increased. The first quantitative data on the species in Italy, coming from two areas (one in the Carnic Alps, the other in the Julian Alps), show high variability among the months in which it was censused. Although impacts and threats are generally considered to be not very serious, there are factors that can affect (in some cases having done so already) even populations living in apparently safe sites, e.g. interventions in environments deriving from human activity, construction works and meteorological events caused by climate change.
Razzetti, E & Bonini, L. & Andreone, F. (2001) -
Common Italian names of the amphibians and reptiles present
in the geographic Italy are proposed. We analysed a large set of
bibliography in order to assess a list of Italian common names
used and we proposed the utilization of a single name, also in
view of the forthcoming publication of the distribution atlas of the
Italian herpetofauna. The criteria used to assign such names follow
in general the current use, the possibility to clearly distinguish
similar species, geographic and linguistic parameters.
Reboredo-Fernández, A. & Ares-Mazás, E. & Galán, P. & Caccio, S.M. & Gómez-Couso (2017) -
Giardia duodenalis is a zoonotic parasite that infects the gut of a wide range of vertebrates, including numerous wildlife species. However, little is known about this protozoan parasite in reptiles. Fecal samples from 31 wild lizards were collected in Galicia (northwest Spain) and screened for the presence of Giardia by PCR amplification and sequencing of the ITS1-5.8S-ITS2 region in the ribosomal unit. This allowed detection of the parasite in 5 samples (16.1%), and enabled identification of G. duodenalis assemblage A2 in two samples of Iberian rock lizard (Iberolacerta monticola), G. duodenalis assemblage B in other two samples of I. monticola, and G. duodenalis assemblage E in one sample of Bocage`s wall lizard (Podarcis bocagei). The results obtained after PCR amplification and sequencing of the SSU-rDNA gene confirmed the presence of G. duodenalis assemblage A in two samples of I. monticola. This is the first report of G. duodenalis in free-living lizards, although further studies are needed to distinguish between actual infection and mechanical dissemination of cysts. The detection of zoonotic and livestock-specific assemblages of G. duodenalis demonstrates the wide environmental contamination by this parasite, possibly due to human activities.
Recio, P. & Rodriguez, G. & Ruiz, F. & Martin, J. (2018) -
Remón, N. & Galán, P. & Naveira, H. (2012) -
The fragmentation and destruction of natural habitats by human intervention is producing a continuous and inexorable reduction of the size of populations in multitude of species all over the world. Small and isolated populations face higher extinction risks, due to demographic and environmental stochasticity, and also because of several genetic threats, among which inbreeding is considered the most important one. For many of these species, the extinction of a population is an irreversible event, so that determining the immediate importance of these risk factors and understanding their interactions is crucial for conservation plans. Iberolacerta monticola is a small lacertid endemic to the northwestern Iberian Peninsula, distributed mainly across moderate/high altitude mountainous regions. Some populations are found nearly at sea-level, though, in fluvial valleys with relict Atlantic forests, in the severely fragmented western part of its range. One of them has been dramatically reduced over the last 30 years, and presently is on the brink of extinction. Using microsatellite nuclear markers, we obtained different measures of genetic variation at this site, together with demographic and breeding data. Both the level of heterozygosity and the number of alleles per locus indicate that the level of variation in this population is comparatively high, and the average inbreeding coefficient is very low. Individuals appear healthy and long-lived, and are related by a few different lines of descent. These findings are discussed in the context of current theories and experimental evidence of associative overdominance and purging of the genetic load of populations, with special emphasis on the evolutionary potential of recovery of small evolutionary units.
Remón, N. & Galán, P. & Vila, M. & Arribas, O. & Naveira, H. (2013) -
Aim The study of the factors that influence population connectivity and spatial distribution of genetic variation is crucial for understanding speciation and for predicting the effects of landscape modification and habitat fragmentation, which are considered severe threats to global biodiversity. This dual perspective is obtained from analyses of subalpine mountain species, whose present distribution may have been shaped both by cyclical climate changes over ice ages and anthropogenic perturbations of their habitats. Here, we examine the phylogeography, population structure and genetic diversity of the lacertid lizard Iberolacerta monticola, an endemism considered to be facing a high risk of extinction in several populations. Location Northwestern quadrant of the Iberian Peninsula. Methods We analyzed the mtDNA variation at the control region (454 bp) and the cytochrome b (598 bp) loci, as well as at 10 nuclear microsatellite loci from 17 populations throughout the distribution range of the species. Results According to nuclear markers, most sampling sites are defined as distinct, genetically differentiated populations, and many of them show traces of recent bottlenecks. Mitochondrial data identify a relatively old, geographically restricted lineage, and four to six younger geographically vicariant sister clades, whose origin may be traced back to the mid-Pleistocene revolution, with several subclades possibly associated to the mid-Bruhnes transition. Geographic range fragmentation of one of these clades, which includes lowland sites, is very recent, and most likely due to the accelerated loss of Atlantic forests by human intervention. Main Conclusion Altogether, the data fit a “refugia within refugia” model, some lack of pattern uniformity notwithstanding, and suggest that these mountains might be the cradles of new species of Iberolacerta. However, the changes operated during the Holocene severely compromise the long-term survival of those genetic lineages more exposed to the anthropogenic perturbations of their habitats.
Remón, N. & Vila, M. & Galán, P. & Naveira, H. (2008) -
Fourteen polymorphic microsatellite loci are described for the Iberian rock lizard, Iberol-
acerta monticola. Genetic variation in a sample of 20 individuals from Piornedo (north-
western Spain) was quantified both by the number of alleles per locus, which ranged from
six to 13, and by the expected frequency of heterozygotes under random mating (heterozy-
gosity), which ranged from 0.761 to 0.902. Single locus and global exclusion probabilities
were also computed, and indicate a high power of these markers for paternity assignments
and mating system studies of I. monticola. All the analysed loci were also polymorphic in
Iberolacerta galani, but only seven in Zootoca vivipara.
Richard, J. & Lapini, L. (1993) -
Roca, V. (2017) -
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.
Rödder, D. & Schulte, U. (2010) -
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.
Rodriguez-Ruiz, G. (2018) -
Rodriguez-Ruiz, G. & López, P. & Martin, J. (2019) -
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.
Rodriguez-Ruiz, G. & POrtega, J. & Cuervo, J.J. & López, P. & Salvador, A. & Martin, J. (2020) -
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.
Rofo Oróns, V. (2015) -
Reptiles, with their great diversity of sex-determining systems, have long been regarded as a model group for studying the evolution of sex determination and sex chromosomes. They also hold a key phylogenetic position to elucidate the organization and evolution of amniote genomes. This PhD thesis aims to contribute to this understanding by investigating sex chromosomes and karyotype evolution in lacertid lizards, with a focus on rock lizard species (genus Iberolacerta) endemic of the Iberian Peninsula. Firstly, we applied classical and molecular cytogenetic methods to identify and characterize previously unknown ZW sex chromosomes in the species I. monticola. Secondly, we developed whole-chromosome paints from I. monticola to detect chromosomal rearragements and test the homology of sex chromosomes among closely related lacertid species. These results revealed a high degree of karyotype conservation, but a rapid and independent differentiation of sex chromosomes, and even a putative cryptic event of sex chromosome turnover. Finally, we explored the mode of evolution of two satellite DNA families shared by all eight Iberolacerta species. Both satellite DNAs showed complex and disparate evolutionary patterns, and a highly dynamic behaviour which may be correlated with chromosomal rearragements and karyotype diversification in this genus.
Rojo, V. & Giovannotti, M. & Naveira, H. & Nisi Cerioni, P.& González-Tizón, A.M. & Caputo Barucchi, V. & Galán, P. & Olmo, E. & Martinez-Lage, A. (2013) -
Rock lizards of the genus Iberolacerta constitute a promising model to examine the process of sex chromosome evolution, as these closely related taxa exhibit remarkable diversity in the degree of sex chromosome differentiation with no clear phylogenetic segregation, ranging from cryptic to highly heteromorphic ZW chromosomes and even multiple chromosome systems (Z1Z1Z2Z2/Z1Z2W). To gain a deeper insight into the patterns of karyotype and sex chromosome evolution, we performed a cytogenetic analysis based on conventional staining, banding techniques and fluorescence in situ hybridization in the species I. monticola, for which previous cytogenetic investigations did not detect differentiated sex chromosomes. The karyotype is composed of 2n = 36 acrocentric chromosomes. NORs and the major ribosomal genes were located in the subtelomeric region of chromosome pair 6. Hybridization signals of the telomeric sequences (TTAGGG)n were visualized at the telomeres of all chromosomes and interstitially in 5 chromosome pairs. C-banding showed constitutive heterochromatin at the centromeres of all chromosomes, as well as clear pericentromeric and light telomeric C-bands in several chromosome pairs. These results highlight some chromosomal markers which can be useful to identify species-specific diagnostic characters, although they may not accurately reflect the phylogenetic relationships among the taxa. In addition, C-banding revealed the presence of a heteromorphic ZW sex chromosome pair, where W is smaller than Z and almost completely heterochromatic. This finding sheds light on sex chromosome evolution in the genus Iberolacerta and suggests that further comparative cytogenetic analyses are needed to understand the processes underlying the origin, differentiation and plasticity of sex chromosome systems in lacertid lizards.
Rojo, V. & Martinez-Lage, A. & Giovannotti, M. & González-Tizón, A.M. & Ceroni, P.N. & Caputo Barucchi, V. & Galán, P. & Olmo, E. & Naveira, H. (2015) -
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.
Rovatsos, M. & Kratochvíl, L. (2017) -
While the stability of sex chromosomes is widely accepted in viviparous mammals and birds, ectothermic vertebrates are still largely viewed as having frequent turnovers in sex determining systems. Frequent changes in sex determining systems in ectotherms could be problematic for field ecological studies as well as for breeding programs, as molecular sexing across a phylogenetically widespread spectrum of ectothermic vertebrates would not be possible. However, we recently documented that sex determining systems in three important reptile lineages (caenophidian snakes, iguanas, and lacertid lizards) are in fact highly conserved. 2.We applied a new molecular procedure to identify sex within each of these three lineages (encompassing altogether around 4,000 species, i.e. nearly 50% of the recent species of reptiles). This technique uses quantitative PCR (qPCR) to compare copy numbers of genes specific for their respective Z (in caenophidian snakes and lacertids) and X (in iguanas) chromosomes between male and female genomes. The DNA samples required can be collected relatively non-invasively. Unlike molecular sexing based on repetitive elements, this technique can be easily applied to previously unstudied species of these lineages, as the number of copies of protein coding genes linked to the differentiated sex chromosomes is evolutionary highly conserved in each. 3.We suggest that qPCR-based molecular sexing using the comparison of gene copy number is a practical choice for non-model species of caenophidian snakes, iguanas and lacertids. Morever, it should also soon be available for other reptile lineages with differentiated sex chromosomes.
Rovatsos, M. & Kratochvíl, L. & Altmanová, M. & Johnson Pokorná, M. (2015) -
Telomeres are nucleoprotein complexes protecting the physical ends of linear eukaryotic chromosomes and therefore helping to ensure their stability and integrity. Additionally, telomeric sequences can be localized in non-terminal regions of chromosomes, forming so-called interstitial telomeric sequences (ITSs). ITSs are traditionally considered to be relics of chromosomal rearrangements and thus very informative in the reconstruction of the evolutionary history of karyotype formation. We examined the distribution of the telomeric motifs (TTAGGG)n using fluorescence in situ hybridization (FISH) in 30 species, representing 17 families of squamate reptiles, and compared them with the collected data from another 38 species from literature. Out of the 68 squamate species analyzed, 35 possess ITSs in pericentromeric regions, centromeric regions and/or within chromosome arms. We conclude that the occurrence of ITSs is rather common in squamates, despite their generally conserved karyotypes, suggesting frequent and independent cryptic chromosomal rearrangements in this vertebrate group.
Rúa, M. & Galán, P. (2003) -
We studied the reproductive ecology of a population of the lacertid lizard Lacerta
monticola at a lowland location in A Coruña (NW Spain) from 1997 to 2002. The timing of the
reproductive cycle was examined based on mark-recapture records of individual lizards in the eld.
The characteristics of the eggs and hatchlings were obtained from clutches laid in the laboratory by
pregnant females that were temporarily removed from the study area. Our results indicate that mating
took place between late March and July. The smallest female with signs of sexualmaturity had a snoutvent
length (SVL) of 52.4 mm and the smallest male was 50.7 mm. Sexual maturity was attained at
the age of 2 years by 41% of the individuals, while the remaining 59% of the lizards matured at the
age of 3 years. The laying period occurred between June and the beginning of August considering all
years. About 61% of the reproductive females produced a single clutch annually, while 39% of the
females produced two clutches per year. Females that produced two clutches were generally larger
(average SVL D 70.9 mm) than those that laid a single annual clutch (average SVL D 62.6 mm). The
mean clutch size was 6.4 eggs (range 4-9). Both clutch size and clutch mass increased signi cantly
with female SVL. The mean egg mass in a clutch decreased signi cantly with clutch size. Hatching
occurred between August and September. Hatching success in the laboratory was 71.7%. The mean
SVL was higher in female hatchlings than in males.
Rucner, D. & Rucner, R. (1971) -
Salvador, A. (1974) -
Salvador, A. (1984) -
Salvador, A. & Diaz, J.A. & Veiga, J.P. & Bloor, P. & Brown, R.P. (2008) -
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.
Sauer, F. (1989) -
Schreiber, E. (1912) -
Schreiber, E. (1913) -
Serrano, F. & Diaz-Ricaurte, J.C. (2019) -
Serre Collet (2018) -
Serre Collet, F. (2018) -
Siliceo, I. & Díaz, J.A. (2010) -
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.
Sillero, N. & Carretero, M.A. & Goncalves, J.A. & Kaliontzopoulou, A. & Marcal, A. (2010) -
Sillero, N. & Gomes, V. & Zagar, A. (2014) -
Silva Mendes, J. da (2018) -
The order Squamata is a species-rich group of reptiles including the groups Sauria, Serpentes and Amphisbaenia. Several members of these groups have been extensively used as model organisms for a great variety of studies of different fields including ecology, behaviour and medicine. Such studies require a well-established phylogenetic framework to trace the evolution and diversity of the studied traits in the squamate tree of life. However, the accuracy of phylogenetic inference in Squamata is unbalanced: while the position of the main squamate groups and high level relationships have been successfully and consistently estimated across phylogenetic studies, the phylogenetic relationships of some groups have provided inconsistent results over the years. The accurate estimation of relationships between these groups might have been hindered by methodological artefacts of the phylogenetic inference due to limited data and analytical tools.
For the last couple of decades, and even presently, phylogenetic inference has relied heavily on mitochondrial DNA as the main molecular marker as a result of its advantages, such as the lack of recombination, easy amplification and high evolutionary rate. While its use is based on sound reasons, some drawbacks and limitations have been highlighted, such as the overall small size, and the fact that all genes are linked thus it represents a single locus. To overcome this, the addition of nuclear DNA complements the use of mitochondrial DNA in phylogenetic inference. Within the nuclear DNA, some markers may present so little variation that it is insufficient to recover phylogenetic relationships, yet slow-evolving nuclear genes have been the most widely used markers in phylogenetic studies. On the other hand, it has been shown that the use of highly informative fast evolving nuclear genes, in combination with mitochondrial DNA, can provide resolution at different parts and depths of the evolutionary history of the species.
Phylogenetic analyses of multiple loci have historically been based on the concatenation approach, .i.e. the combination of the sequences into a single alignment that is then analysed to represent the species tree. This method, however, relies on the assumption that the most commonly occurring gene tree is equivalent to the true species tree of any taxa, which may not be true in all cases. Another limitation of the concatenation approach is the elevated percentage of missing data that it sometimes entails, especially in large taxon sets, meaning that the sequence representation is The order Squamata is a species-rich group of reptiles including the groups Sauria, Serpentes and Amphisbaenia. Several members of these groups have been extensively used as model organisms for a great variety of studies of different fields including ecology, behaviour and medicine. Such studies require a well-established phylogenetic framework to trace the evolution and diversity of the studied traits in the squamate tree of life. However, the accuracy of phylogenetic inference in Squamata is unbalanced: while the position of the main squamate groups and high level relationships have been successfully and consistently estimated across phylogenetic studies, the phylogenetic relationships of some groups have provided inconsistent results over the years. The accurate estimation of relationships between these groups might have been hindered by methodological artefacts of the phylogenetic inference due to limited data and analytical tools.
For the last couple of decades, and even presently, phylogenetic inference has relied heavily on mitochondrial DNA as the main molecular marker as a result of its advantages, such as the lack of recombination, easy amplification and high evolutionary rate. While its use is based on sound reasons, some drawbacks and limitations have been highlighted, such as the overall small size, and the fact that all genes are linked thus it represents a single locus. To overcome this, the addition of nuclear DNA complements the use of mitochondrial DNA in phylogenetic inference. Within the nuclear DNA, some markers may present so little variation that it is insufficient to recover phylogenetic relationships, yet slow-evolving nuclear genes have been the most widely used markers in phylogenetic studies. On the other hand, it has been shown that the use of highly informative fast evolving nuclear genes, in combination with mitochondrial DNA, can provide resolution at different parts and depths of the evolutionary history of the species.
Phylogenetic analyses of multiple loci have historically been based on the concatenation approach, .i.e. the combination of the sequences into a single alignment that is then analysed to represent the species tree. This method, however, relies on the assumption that the most commonly occurring gene tree is equivalent to the true species tree of any taxa, which may not be true in all cases. Another limitation of the concatenation approach is the elevated percentage of missing data that it sometimes entails, especially in large taxon sets, meaning that the sequence representation is Miocene and supported a scenario of east-to-west diversification. The little morphological and phylogenetic evidence for the distinctiveness between Rhinechis and Zamenis supports a classification lumping which better reflects their evolutionary history and, based on the priority rule, R. scalaris is moved into the genus Zamenis and designated as Zamenis scalaris comb. nov.
Within the Gallotiinae subfamily, the combined use of fast nuclear markers and the species tree was fruitful for the inference of a robust phylogeny of the genus Psammodromus, distributed in the Iberian Peninsula and North Africa. In this study we analysed for the first time the six Psammodromus species to infer the association between the main cladogenetic events within this genus and the complex biogeographic dynamics across the Strait of Gibraltar. The inferred phylogeographical history suggests that Psammodromus probably originated in Iberia. The African species and the African lineage of P. algirus were a result of two over-sea dispersal events towards Africa that occurred 10 Ma and 1.5 Ma, while continental vicariance events might have shaped the diversification of the species within Iberia and Africa. These results, combined with previous literature, provide compelling evidence that major biotic exchanges occurred across the Strait of Gibraltar well before or long after the land connection during the Messinian Salinity Crisis (5.9 – 5.33 Ma). These findings suggest caution in the application of the relatively short event of the opening of the Strait of Gibraltar at the end of the Messinian Salinity Crisis as a cause for divergence in molecular clock calibrations, which is a common approach in literature.
At the species level, the use of fast evolving nuclear markers proved to be fruitful to understand the geographical pattern of phyletic diversification within the Omanosaura lizards belonging to the Eremiadini tribe, endemic to the Hajar Mountains in the Arabian Peninsula. Multilocus phylogenetic analyses recovered two highly divergent lineages within O. cyanura which are geographically associated to the northernmost and to the south and eastern regions of the Hajar Mountains. These lineages are reciprocally monophyletic at both mitochondrial and nuclear loci suggesting a long history of independent evolution and the need of a comprehensive taxonomic assessment. Omanosaura cyanura represents an additional case of cryptic diversity in the north Hajar Mountains, an area that has been demonstrated to hide high levels of genetic diversity in other reptile groups.
Overall, the work developed in this dissertation has demonstrated that the inclusion of fast evolving nuclear genes and the coalescent species tree approach was relevant for resolving challenging phylogenetic questions on selected squamate groups. This approach allowed the recovery of new clades and provided support for old and recent relationships. These methods have further allowed the comparison between contrasting phylogenetic hypotheses on the Lacertini and Colubrinae, demonstrating that the inference based on the supermatrix approach used by many recent studies may provide high support for incorrect nodes. Future research directions include (i) the application of the phylogenomic approach based on Next Generation Sequence data to further resolve basal polytomy within the Lacertini and Colubrinae radiations; (ii) a taxonomical assessment of Omanosaura; (iii) a re-evaluation of molecular clock calibrations based on the end of the Messinian Salinity Crisis as general cause for divergence, in order to account for biotic exchanges by over-sea dispersal across the Mediterranean well before or long after the land connection during this period, as suggested by emerging literature and the study case on Psammodromus.
Sindaco, R. & Paggetti, E. (2011) -
Sochurek, E. (1955) -
Sochurek, E. (1986) -
Sociedade Galega de Historia Natural (2019) -
Speybroeck, J. (2004) -
Speybroeck, J. (2005) -
Speybroeck, J. (2006) -
Speybroeck, J. (2010) -
Speybroeck, J. (2012) -
Speybroeck, J. & Beukema, W. & Crochet, P.-A. (2010) -
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.
Speybroeck, J. & Beukema, W. & Dufresnes, C. & Fritz, U. & Jablonski, D. & Lymberakis, P. & Martinez-Solano, I. & Razzettis, E. & Vamberger, M. & Vences, M. & Vörös, J. & Crochet, P.-A. (2020) -
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).
Speybroek, J. (2010) -
Steinicke, H. & Henle, K. & Gruttke, H. (2002) -
Steinicke, H. & Henle, K. & Gruttke, H. (2002) -
Strijbosch, H. (1986) -
Szczerbak, N.N. (1982) -
Elements of the herpetofauna are in the author`s opinion of great importance for the zoogeographical division of the Palaearctic region. It is suggested that this region be divided into the following four subregions: 1/ arctic, 2/ boreal Euro-Siberian, 3/ bo- real Himalayan-Manchurian and 4/ Mediterranean-Central-Asiatic.
Szirovicza, L. & López, P. & Kopena, R. & Benkö, M. & Martin, J. & Penzes, J.J. (2016) -
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.
Taddei, A. (1950) -
Thirion, J.M. & Vollette, J. & Plisson, C. & André, C. & Lafitte, J. & Sourp, E. (2018) -
Monitoring of the population of the Pyrenean rock lizard Iberolacerta bonnali (Lantz, 1927) in the “Val d’Arrious” (Ossau Valley). A monitoring of the population of the Pyrenean rock lizard Iberolacerta bonnali in the Val d’Arrious (Ossau Valley) was established in the Pyrenees National Park. This monitoring allowed mapping the distribution of the Pyrenean rock lizard’s as well as the contact zone with the common wall lizard Podarcis muralis, from the altitude of 2.010 m a.s.l upwards. The habitat of the Pyrenean rock lizard is restricted to the most thermophilous exposures, including Festuca eskia meadows, screes areas, siliceous slabs, and outcrops along with damp boulders and screes where the snow remains, along the stream of Arrious. A capture-mark-recapture study estimated the population size of the Pyrenean Rock Lizard’s population size at 69 individuals (54 - 99 individuals at 95 %), for 2.458 m², i.e. a density of 281 lizards per hectare. The study of the Pyrenean rock lizard of Arrious allowed to make a first inventory of this species and to validate a monitoring method. Other sites will be studied following this protocol to establish a network of sites for long-term monitoring.
Tiedemann, F. (1987) -
Tiedemann, F. (1992) -
The story of discovery of Austrian populations of Horvath`s Rock Lizard (Lacerta horvathi MÉHELY, 1904) is briefly outlined. A chart of all locality records hitherto known in Austria (Cannthia) is provided, including first records from the Karawanken Mts. The fruitless attempts to confirm Bavarian records of this species and to ascertain its presence in the Tyrol are reported.
Tiedemann, F. (1997) -
Tiedemann, F. & Grillitsch, H. (1999) -
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.
Tvrtković, N. (1984) -
Tvrtković, N. & Kletečki, E. (1998) -
Tvrtković, N. & Veen, P. (2006) -
Urosevic, A. & Ljubisavljevic, K. & Ivanovic, A. (2013) -
We explored the ontogenetic dynamics of the morphological and allometric disparity in the cranium shapes of twelve lacertid lizard species. The analysed species (Darevskia praticola, Dinarolacerta mosorensis, Iberolacerta horvathi, Lacerta agilis, L. trilineata, L. viridis, Podarcis erhardii, P. melisellensis, P. muralis, P. sicula, P. taurica and Zootoca vivipara) can be classified into different ecomorphs: terrestrial lizards that inhabit vegetated habitats (habitats with lush or sparse vegetation), saxicolous and shrub-climbing lizards. We observed that there was an overall increase in the morphological disparity (MD) during the ontogeny of the lacertid lizards. The ventral cranium, which is involved in the mechanics of jaw movement and feeding, showed higher levels of MD, an ontogenetic shift in the morphospace planes and more variable allometric patterns than more conserved dorsal crania. With respect to ecology, the allometric trajectories of the shrub-climbing species tended to cluster together, whereas the allometric trajectories of the saxicolous species were highly dispersed. Our results indicate that the ontogenetic patterns of morphological and allometric disparity in the lacertid lizards are modified by ecology and functional constraints and that the identical mechanisms that lead to intraspecific morphological variation also produce morphological divergence at higher taxonomic levels.
We used Procrustes-based geometric morphometrics to explore morphological variability in dorsal and ventral cranium shape of twelve lacertid lizard species from the Balkan Peninsula (), at the juvenile (neonatal) and at the adult stage (females and males separately). These species differed in phylogenetic relatedness, overall size and habitat preference (terrestrial in overgrown habitats, terrestrial in sparsely vegetated habitats, saxicolous and semiarboreal). General pattern of shape variation appeared to be preserved throughout ontogeny, especially for the dorsal cranium - adult female and male morphospaces corresponded to the neonate morphospace. The inspection of morphospaces showed that the general pattern of shape variability was along the gradient from the smaller to the species with larger body size. Along this gradient, species clustered according to habitat preference. The main difference between neonates and adults was the position of semiarboreal species, which completely separated from the rest of the species at the adult stage. The overall morphological disparity (MD) increased during the course of ontogeny (from neonates to adult females and males). Ventral cranium, with its structures involved in mechanics of jaw movement and feeding, showed greater increase in MD, as well as the shift in the morphospace hyperplanes. On the generic level, Lacerta spp. showed significantly higher MD than Podarcis spp. In contrast to other ecological groups (terrestrial and semiarboreal) saxicolous lizards showed a tendency of decreasing shape disparity during ontogeny. The species from saxicolous group were phylogenetically heterogeneous but morphologically convergent due to the specific habitat constraints, and their distinctive MD pattern could be achieved by different allometric paths. The patterns of shape variation and MD were modified by ecology, functional constraints and different ontogenetic trajectories. Further studies on the ecomorphology, allometric diversity and morphological integration, as well as the reassessment of the problematic lacertid phylogeny are needed in order to shed more light on the complex relationships among morphology, ecology and phylogeny in lacertid lizards.
Urosevic, A. & Ljubisavljevic, K. & Ivanovic, A. (2018) -
Different factors and processes that produce phenotypic variation at the individual, population, or interspecific level can influence or alter the covariance structure among morphological traits. Therefore, studies of the patterns of integration and modularity at multiple levels—static, ontogenetic, and evolutionary, can provide invaluable data on underlying factors and processes that structured morphological variation, directed, or constrained evolutionary changes. Our dataset, consisting of cranium shape data for 14 lizard species from the family Lacertidae, with substantial samples of hatchlings and adults along with their inferred evolutionary relationships, enabled us to assess modularity and morphological integration at all three levels. Five, not mutually exclusive modularity hypotheses of lizard cranium, were tested, and the effects of allometry on intensity and the pattern of integration and modularity were estimated. We used geometric morphometrics to extract symmetric and asymmetric, as well as allometric and nonallometric, components of shape variation. At the static level, firm confirmation of cranial modularity was found for hypotheses which separate anterior and posterior functional compartments of the skull. At the ontogenetic level, two alternative hypotheses (the “anteroposterior” and “neurodermatocranial” hypotheses) of ventral cranial modularity were confirmed. At the evolutionary level, the “neurodermatocranial” hypothesis was confirmed for the ventral cranium, which is in accordance with the pattern observed at the ontogenetic level. The observed pattern of static modularity could be driven by functional demands and can be regarded as adaptive. Ontogenetic modularity and evolutionary modularity show the same developmental origin, indicating conservatism of modularity patterns driven by developmental constraints.
Vacher, J.-P. & Geniez, M. (2010) -
Vasilopoulou-Kampitsi, M. & Goyens, J. & Damme, R. van & Aerts, P. (2019) -
Shape variation in the vestibular system is often linked to microhabitat structure and locomotor performance. Highly circular and orthogonal semicircular canal pairs are linked to higher motion sensitivity. Here, we use 3D geometric morphometrics to investigate shape variation in the vestibular system within lacertid lizards and its relationship to balance control. We found that lacertids living in complex microhabitats possess narrow but longer vestibular systems, an S-shaped anterior canal, a straightened lateral canal and a short common crus. However, lacertids specialized for simple microhabitats (open areas) possess wider but shorter vestibular systems, more circular anterior and lateral canals, and a longer common crus. Contrary to our expectations, species living in simple microhabitats possess more anatomical adaptations that enhance the sensitivity of their vestibular system. This suggests that species inhabiting open areas may benefit from increased sensitivity given that they are potentially more visibile to predators and have lower shelter availability. Finally, the wider shape of the vestibular system of the open area species may be linked to a wider and potentially flattened skull, which may be related to sand-diving or prey hardness.
Veith, G. (1991) -
Vences, M. & Rey, J. & Puente, M. & Miramontes, C. & Dominguez, M. (1998) -
Vences, M. & Thiesmeier, B. & Glaw, F. (1992) -
Venchi, A. & Vignoli, L. & Luiselli, L. (2011) -
Verwaijen, D. & Van Damme, R. (2007) -
Evolutionary changes in foraging style are often believed to require concurrent changes in a complex suite of morphological, physiological, behavioural and life-history traits. In lizards, species from families with a predominantly sit-and-wait foraging style tend to be more stocky and robust, with larger heads and mouths than species belonging to actively foraging families. Here, we test whether morphology and foraging behaviour show similar patterns of association within the family Lacertidae. We also examine the association of bite force abilities with morphology and foraging behaviour. Lacertid lizards exhibit considerable interspecific variation in foraging indices, and we found some evidence for a covariation between foraging style and body shape. However, the observed relationships are not always in line with the predictions. Also, the significance of the relationships varies with the evolutionary model used. Our results challenge the idea that foraging style is evolutionarily conservative and invariably associated with particular morphologies. It appears that the flexibility of foraging mode and its morphological correlates varies among lizard taxa.
Verwaijen, D. & Van Damme, R. (2008) -
Apart from certain species of the African clade, the lizard family Lacertidae has generally been
described as consisting of active foragers. We made quantitative field observations of 14 species of lacertid
lizards, mainly belonging to the more basal Eurasian clade. Our data show that sit-and-wait foraging is much
more widespread in Lacertidae than previously alleged. We also investigated the influence of weather and
times of day on foraging activity levels and conclude that for comparative purposes observations should be
restricted to circumstances that are optimal for activity. We did not find sex differences in foraging behavior.
Foraging strategy is often considered to play a central role in moulding diverse aspects of an animal`s general biology. Active foragers should have greater locomotor endurance, allowing high movement activity rates, while sit-and-waiting foragers may be better adapted to sprinting, allowing catching prey by a quick attack from an ambush site, and going with specific predator escape tactics. In this study we investigate these predicted patterns in a set of lacertid lizard species. There is considerable variation in foraging activity within Lacertidae, which allows the close investigation of the co-evolution of the traits considered. We found a tendency of positive correlation of foraging measures (PTM, percentage of time moving; MPM, number of movements per minute) with laboratory measured endurance capacity. However, the relationship of foraging measures with maximal sprint speed remains less clear. MPM correlates negatively with maximal sprint speed, but PTM does not. When sprint speed was corrected for body size, no correlations were found at all.
Space usage by animals may be influenced by a range of factors. In this study we investigate whether foraging behaviour affects the home range size of lizards. Two distinct tactics of foraging have been recognized in predators: sit-and-wait foraging (SW) and active foraging (AF). Foraging activity level of a data set of lizard species, mainly compiled from literature, is compared with their home range sizes. Two opposite predictions can be made about foraging in connection with home range area: on the one hand, SW species may exhibit larger home ranges due to their mating system; on the other hand, AF species have higher metabolic energy and thus food requirements and can be expected to have larger home ranges that have to yield this food. This study shows that percentage of the time moving (as an index of foraging mode) correlates positively with home range, even after correcting for body mass, and these patterns remain when phylogenetic relationships are taken into account. We thus conclude that home range areas parallel activity levels in lizards.
Villa, A. & Delfino, M. (2019) -
The fossil record provides evidence of a long evolutionary history of European lizards. Since fossil lizards are regularly represented by bone remains, the knowledge of the origins of extant taxa and their distribution in time and space is hindered by the fact that their comparative osteology is not yet completely and adequately known. In spite of a rising interest in this topic since the end of the 20th century, a gap in our knowledge is still evident. We here report the first broad-scale comparative osteological analysis of the skulls of extant European lizards, highlighting significant differences that can be used in identification. This comparative study, including as many European species as possible, leads to the creation of a detailed diagnostic key for each single bone. Also, our data significantly improve the recognizability of extant European non-snake squamates, with 54% of the current diversity to be recognized based on the new results contra the previously estimated 31%. This recognizability is expected to further increase in the future, with new studies focusing on species that are either missing or poorly represented here, or applying promising advanced methodologies.
Vogrin, M. & Böhme, W. & Crochet, P.-A. & Nettmann, H.K. & Sindaco, R. & Romano, A. (2009) -
Voort, J. van der (2004) -
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.
Wagner, N. & Mingo, V. & Schulte, U. & Lötters, S. (2015) -
Environmental contamination is supposed to be a reason for population declines in reptiles. Especially intensification and expansion of agriculture are leading to increased pesticide exposure risks for wildlife. In the European Union, Special Areas of Conservation (SACs) have been established for the conservation of taxa listed in Annex II of the Habitats Directive. In the SACs, agricultural land use is legal. Therefore, we conducted a risk evaluation of pesticide exposure for Annex II reptiles by calculating proportions of land use with regular pesticide applications within SACs. Using three evaluation factors (occurrence probability, physiology, life-history aspects), a species-specific risk index was created. Nearly half of the species at above-average risk by pesticide use are globally threatened with extinction (IUCN Red List of Threatened Species). About 30% of their SACs are agriculturally used and one priority subspecies of the Habitats Directive is at highest risk (Vipera ursinii rakosiensis). Also, all evaluated fresh-water and land-dwelling turtle species are at high risk. National variation in agricultural land use in the SACs was observed. Species at above-average risk are mainly distributed in the Mediterranean and Pannonian/Continental biogeographical regions of Europe. Conservation status according to the IUCN Red List of Threatened Species as well as national differences among the member states argue for the inclusion of pesticide risk assessments in site-specific management plans for SACs to avoid regional loss of reptilian biodiversity.
Werner, F. (1906) -
Wettstein, O. (1928) -
Zagar, A. (2008) -
Zagar, A. (2010) -
Zagar, A. (2014) -
Zagar, A. (2016) -
The study reports on the distribution and habitat use of two lizard species in the Kočevsko region: Horvath’s rock lizard and common wall lizard. Extensive sampling across an altitudinal span of 200 to 1,100 m a.s.l. in the study area revealed 62 localities with populations of both or either species. At 11 of these localities (18%) species occurred in syntopy, at 42 locations (68%) only common wall lizards were found, while at 9 locations (14%) only Horvath’s rock lizards were recorded. Both species occurred across the entire altitudinal span but exhibited an opposite pattern of relative abundances and frequencies, which increased with increasing altitude in Horvath’s rock lizard and with decreasing altitude in common wall lizard. The habitat use of common wall lizard was more general (it was found in seven habitat types) than Horvath’s rock lizard that was registered only in three habitat types with rocks.
The understanding of mechanisms of interspecific competition between ecologically similar sympatric species is often lacking from studies predicting the impact of future climate changes. However biotic interactions are known to be key factors influencing the community structure. We studied interspecific competitive mechanisms in two species: the Horvath`s rock lizard (Iberolacerta horvathi [Méhely, 1904]) and the common wall lizard (Podarcis muralis [Laurenti, 1768]). At first we examined the potential for competition between studied species by comparing their realised niches in the sympatric area and their morphological, physiological and ecological traits (species` fundamental ecological niches). Altitude contributed the most to the segregation pattern in their distribution. Both species have similar general habitat use and were morphologically similar, except for the head height, which might have reprecursions in segregation in the use of refuge sites. Species were similar in the mean of preferred body temperatures and metabolic rates, but differed in the precision of thermoregulation and the potential metabolic activity. Thus, I. horvathi might have eco-physiological competitive advantages over P. muralis in thermally restrictive (cold) environments. Furthermore, species were found to compete for limited resource (sun-exposed areas) via direct agonistic interactions, indicating an asymmetric interspecific interaction (P. muralis showed to be in adavnatge over I. horvathi). We have also assesed the role of common predators as mediators on the interaction between lizards as prey species. We suggest that “a more cautious” anti-predator behavioural response of I. horvathi might provide direct benefits (higher survival rates) but only in populations where predation pressure was high, otherwise it carries substantial time and thermoregulatory costs. Overall results of the dissertation improved our understanding of interspecific competition between studied sympatric species. Such information may also be used for improving results of modelling predictions of the effect of future climate changes on the distribution of these protected species.
Zagar, A. & Bitenc, K. & Vrezec, A. & Carretero, M.A. (2015) -
Prey response to different predators is complex and can include diverse antipredatory strategies. In syntopic populations of competing species common predators can play a mediator role thus influence the interaction out-come of prey-species in an indirect way. We studied differences in antipredator response in two competing lizards in syntopy in a multipredator environment. Studied prey species, Iberolacerta horvathi and Podarcis muralis, are likely to compete in syntopic populations limited in size and have similar morphology and ecology but exhibit fine-scale ecophysiological differences. Taking into account interspecific differences in ecophysiology we expected that I. horvathi as a more precise thermoregulator would be less prone to use refuges that represent a thermal cost (are colder than outside). The pattern we found was the opposite of our expectations; I. horvathi escaped at greater distances and remained in the refuge for longer before re-emerging than P. muralis. Second part of the study revealed that both species were able to recognize chemical cues of predator snakes (represented as higher tongue flick rates in the presence of scents of predator snakes in comparison to control). Behavioral responses, which are linked with stressful situations connected to saurophagous snakes, were more frequent and variable in I. horvathi. Overall, antipredator responses seem to be more pronounced in /. horvathi than in P. muralis. While this `more cautious` attitude of I. horvathi should provide higher short-term benefits (higher survival rates), this would hold true only in populations where predation pressure is high. Otherwise it carries substantial time and thermoregulatory costs. For I. horvathi, costs of refuge use should be even higher due to narrower dimensions of their ecophysiological fundamental niche. Such divergences in antipredator behavior are expected to shape the relationships between both species in syntopic populations modulated by common predation pressure and habitat structure.
Zagar, A. & Carretero, M.A. (2014) -
Zagar, A. & Carretero, M.A. & Krofel, M. & Luznik, M. & Podnar, M. & Tvrtkovic, N. (2014) -
The article presents results from a reptile survey conducted between 14 and 16 June 2013 in a montane and subalpine belt of the western part of Dinara Mountain in Croatia. The most interesting result is the southernmost finding of Horvath’s Rock lizard (Iberolacerta horvathi). In the survey area we also recorded eight other reptile species: Anguis fragilis, Lacerta agilis, L. viridis sensu lato, Podarcis muralis, P. melissellensis, Coronella austriaca, Zamenis longissimus, and Vipera ammodytes.
Zagar, A. & Carretero, M.A. & Osojnik, N. & Sillero, N. & Vrezec, A. (2015) -
Solar radiation is in theory an unlimited resource on Earth, but can be locally limited. Heliothermic organisms use solar radiation to elevate their body temperatures, leading them to compete for sun-exposed areas, where interference for limited resource can occur. In coexisting lizard species, interference for basking sites could promote and direct interspecific interactions, which can be related to broader geographic species distribution patterns. We compared two competing lizard species (Podarcis muralis and Iberolacerta horvathi) that occupy sun-exposed gaps in a forested landscape in the Northern Dinaric Mountains (southern Europe). Both species exhibit an altitudinal segregation pattern with a high zone of overlap at middle altitudes. Using experimental thermal gradients, we tested for existence of interference competition. We used adult male lizards in three social contexts, alone as a measure of preferred body temperature (T p) and in conspecific or heterospecific pairs as a measure of realized body temperature (T r). I. horvathi achieved lower T r compared to its T p (alone), while T r of P. muralis did not vary between social contexts. The evidence for interference found in I. horvathi could not be explained by a change in their general movement pattern in a thermal gradient. Our results suggest that interference was a result of direct agonistic interactions, assessed by the frequency of approaches and touches. I. horvathi showed higher interference-susceptibility than P. muralis indicating an asymmetric interaction between the two species. Sunlight (as a resource) is an important factor influencing distribution patterns in assemblages of heliothermic ectotherms where sun-exposed basking sites are limited.
Zagar, A. & Carretero, M.A. & Vrezec, A. (2013) -
Lacertids are preyed upon by different predators with different foraging strategies, namely, avian predators attack from above, while terrestrial predators use either approaching on open ground or active searching for prey in their hiding places. When active, lacertids are expected to employ different antipredatory behaviours, either crypsis or early predator detection (visually or chemically) followed by escape to the shelter. Syntopic populations of overall similar lacertid species are likely to share predators. However, their predator avoidance tactics might differ. Using standard techniques, we conducted a field test of the escape-recovery tactics for two morphologically and ecologically similar species: Iberolacerta horvathi and Podarcis muralis, occurring in syntopy in the Northern Dinaric region. Adults of both species intrinsically differed in their escape tactics when approached by the researcher simulating a ground predator attack. Specifically, I. horvathi tended to escape at greater distances and spent more time in the shelter before emerging (longer recovery time) regardless the air, surface or refuge temperatures and habitat type. These divergent antipredator strategies involving different predation risks are expected to affect coexistence of these two lizard species.
Žagar, A. & Carretero, M.A. & Vrezec, A. & Drasler, K. & Kaliontzopoulou, A. (2016) -
Functional performance linked with individual fitness has a major impact on population survival. In sympatric species where the potential for competition is high due to ecological resemblance, variation in performance may play an important role in facilitating species co-existence. Sexual selection also influences morphology. We aimed to examine the potential role of functional morphology and whole-organism performance in species co-existence patterns. We investigated functional traits of two sympatric species, Iberolacerta horvathi and Podarcis muralis. They exhibited a partial segregation pattern, but were found to co-exist in 18% of all populations in the study area. We captured between 24 and 28 females and males of both species to quantify morphological traits and performance (bite-force and speed). Observed variation in functional traits suggests four major mechanisms probably enhancing co-existence: (i) Head shape in connection with bite-force may determine trophic segregation and (ii) may influence success in agonistic social encounters. Males had higher bite forces than females, and P. muralis exerted higher bite forces than I. horvathi of the same sex. Relatively higher but narrower heads were associated with increased biting performance in P. muralis. (iii) Difference in head height may allow spatial segregation in the use of crevices that is potentially linked with different prey, egg-laying sites and predator avoidance. Iberolacerta horvathi had a flatter head than P. muralis. (iv) Size of female trunk-length may promote the relative size of a clutch and positively influence reproductive effort by clutch. Since higher climbing speed was associated to longer limbs and shorter trunks, females (especially I. horvathi) were the slowest climbers. In conclusion, head dimensions and bite force were observed to be ecomorphological traits potentially involved in promoting co-existence between species. We also found a connection between trunk length and climbing performance. However, its implication for species interactions is more difficult to understand, possibly because strong selection pressures for reproduction are involved.
Zagar, A. & Carretero, M.A. & Vrezec, A. & Drasler, K. & Kaliontzopoulou, A. (2017) -
1. We examined intra- and interspecific variation in functional morphology and whole-organism performance in a sympatric lizard species pair, Iberolacerta horvathi and Podarcis muralis, in the area with a high potential for competition. 2. The biggest variation between species was found in two functional traits, bite force and climbing speed, linked with corresponding morphological traits. 3. The species with larger and taller heads, P. muralis, exhibited correspondingly stronger bite forces. The other species exhibited smaller and flatter head. Both traits may potentially promote segregation between species in trophic niche (stronger bites relate to harder prey) and in refuge use (flatter heads allow using narrower crevices, hence, influencing escaping from common predators). Stronger bites and larger heads also provide one species with a dominant position in interspecific agonistic interactions. 4. Females had longer trunks that impacted negatively on climbing speed, which may lower anti-predator escape abilities of the more trunk-dimorphic species, but positively influence reproductive effort. 5. Our results exemplify how the joint examination of morphological and functional traits of ecologically similar and sympatric species can provide a mechanistic background for understanding their coexistence, namely syntopic populations that are frequent in the study area. 6. The identified roles of functional morphology in this system of sympatric rock lizards support the contribution of functional diversification for the complexity of community structure via coexistence.
Zagar, A. & Kos, I. & Vrezec, A. (2013) -
We have surveyed sympatric reptiles in a diverse and preserved montane environment in the Northern Dinaric region (Southern Slovenia) to determine the assemblage structure and assess the patterns of habitat segregation. Altitude and habitat type contributed the most to segregations between the most abundant species. The most similar were species pairs of L. viridis/bilineata and P. muralis, and I. horvathi and V. ammodytes. In snakes, significant segregation patterns have been observed between all species. Among lizards, we have found strong altitudinal segregation between two morphologically and ecologically most similar species; P. muralis was most abundant at low and I. horvathi at higher elevations. This result indicated a potential competitive interaction between these species
Žagar, A. & Osojnik, N. & Carretero, M.A. & Vrezec, A. (2012) -
Podarcis muralis and Iberolacerta horvathi are sympatric, frequently syntopic,
lacertids through the entire range of I. horvathi and very similar in their general
body size and shape, as well as in most ecological traits. We morphologically compared
adults from the area of sympatry using biometric measurements and performed
analyses to investigate their sexual size and shape dimorphism. A total of 34 males
and 24 females of I. horvathi, and 25 males and 23 females of P. muralis, all adult individuals,
were measured. Both species showed sexual size dimorphism with females
being longer (snout-vent length, SVL) than males. After SVL correction (ANCOVA),
head width, length and height and mass showed to be sexually dimorphic in both species.
Males carry relatively wider, longer and higher heads and were heavier than conspecific
females. I. horvathi heads were more flattened than those of P. muralis and P.
muralis were heavier than I. horvathi. Both species displayed the same pattern of sexual
dimorphism regarding body