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Die landgebundenen Wirbeltiere des festländischen Iberiens (Süßwasserfische, Amphibien, Reptilien mit Ausnahme der Meeresschildkröten, Säugetiere mit Ausnahme der Chiroptera, Pinnipedia und Cetacea) bilden, wie im einzelnen gezeigt wird, in ihrer Gesamtheit eine besondere faunistische Einheit: ungefähr die Hälfte der Formen (Spezies oder Subspezies) ist endemisch; das gilt für 73 % der echten Süßwasserfische, 47 % der Amphibien, 36 % der Reptilien und 53 % der Säugetiere. Die geographische Isolierung durch die Pyrenäen und das Mittelmeer ist gewiß die Hauptursache für die Entwicklung einer so eigenartigen Fauna, aber auch andere Faktoren (die sehr verschiedenen Klimate und Biotope innerhalb Iberiens, der im Vergleich mit anderen Gebieten Europas geringere Einfluß der Vereisungen usw. haben sicher ebenfalls eine sehr wichtige Rolle gespielt. Acanthodactylus erythrurus erythrurus, Algyroides hidalgoi, Algyroides marchi marchi, Algyroides marchi niethammeri, Lacerta agilis agilis, Lacerta hispanica hispanica, Lacerta hispanica bocagei, Lacerta hispanica vaucheri, Lacerta lepida lepida, Lacerta lepida nevadensis, Lacerta monticola monticola, Lacerta monticola cantabrica, Lacerta monticola cyreni, Lacerta schreiberi, Lacerta viridis viridis, Lacerta vivipara, Psammodromus algirus algirus, Psammodromus hispanicus hispanicus, Psammodromus hispanicus edwardsianus.

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Geographical body size variation has long interested evolutionary biologists, and a range of mechanisms have been proposed to explain the observed patterns. It is considered to be more puzzling in ectotherms than in endotherms, and integrative approaches are necessary for testing non-exclusive alternative mechanisms. Using lacertid lizards as a model, we adopted an integrative approach, testing different hypotheses for both sexes while incorporating temporal, spatial, and phylogenetic autocorrelation at the individual level. We used data on the Spanish Sand Racer species group from a field survey to disentangle different sources of body size variation through environmental and individual genetic data, while accounting for temporal and spatial autocorrelation. A variation partitioning method was applied to separate independent and shared components of ecology and phylogeny, and estimated their significance. Then, we fed-back our models by controlling for relevant independent components. The pattern was consistent with the geographical Bergmann`s cline and the experimental temperature-size rule: adults were larger at lower temperatures (and/or higher elevations). This result was confirmed with additional multi-year independent data-set derived from the literature. Variation partitioning showed no sex differences in phylogenetic inertia but showed sex differences in the independent component of ecology; primarily due to growth differences. Interestingly, only after controlling for independent components did primary productivity also emerge as an important predictor explaining size variation in both sexes. This study highlights the importance of integrating individual-based genetic information, relevant ecological parameters, and temporal and spatial autocorrelation in sex-specific models to detect potentially important hidden effects. Our individual-based approach devoted to extract and control for independent components was useful to reveal hidden effects linked with alternative non-exclusive hypothesis, such as those of primary productivity. Also, including measurement date allowed disentangling and controlling for short-term temporal autocorrelation reflecting sex-specific growth plasticity.

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I visited southern France to search for Timon lepidus for the third time in 1998. Sites visited included (among others) Maussane, Lac du Salagou and the surroundings of this lake, St. Rémy-de-Provence, the surrounding hills of Le petit Crau and Avignon. I did not find Timon lepidus this time, but did see Natrix maura, Malpolon monspessulanus, Lacerta bilineata, Podarcis muralis and many Psammodromus hispanicus. There were not very many amphibians visible because of the hot weather.

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Algyroides fitzingeri, Algyroides marchi, Algyroides moreoticus, Algyroides nigropunctatus, Lacerta agilis, Lacerta lepida, Lacerta princeps, Lacerta schreiberi, Lacerta strigata, Lacerta trilineata, Lacerta viridis, Lacerta andreanszkyi, Lacerta armeniaca, Lacerta bedriagae, Lacerta brandtii, Lacerta cappadocica, Lacerta caucasica, Lacerta chlorogaster, Lacerta cyanura, Lacerta danfordi, Lacerta derjugini, Lacerta dugesii, Lacerta fraasii, Lacerta graeca, Lacerta horvathi, Lacerta jayakari, Lacerta laevis, Lacerta monticola, Lacerta mosorensis, Lacerta oxycephala, Lacerta parva, Lacerta perspicillata, Lacerta praticola, Lacerta rudis, Lacerta saxicola, Lacerta vivipara, Gallotia atlantica, Gallotia galloti, Gallotia simonyi, Podarcis erhardii, Podarcis filfolensis, Podarcis hispanica, Podarcis lilfordi, Podarcis melisellensis, Podarcis milensis, Podarcis muralis, Podarcis peloponnesiaca, Podarcis pityusensis, Podarcis sicula, Podarcis taurica, Podarcis tiliguerta, Podarcis wagleriana, Psammodromus algirus, Psammodromus blanci, Psammodromus hispanicus, Psammodromus microdactylus.

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The basic anatomy of the lacertid hemipenis (intromittent organ) and methods for its investigation are described. In many members of the Lacertidae, the hemipenis has a structure quite unlike that of other squamate reptiles: the distal lobes of the retracted organ are complexly folded and there is a well-defined supporting structure of dense connective tissue, the armature. This incorporates blood sinuses and has an intramuscular portion embedded in the m. retractor penis magnus and two club-shaped bodies, the clavulae, that support the lobes in the erect organ. Unarmatured hemipenes occur in some lacertids and, like those of other squamates, possess sac-like lobes in the retracted state, but they are singular in having the lobes invested by the m. retractor penis magnus. It is argued that many of these apparently primitive hemipenes are in fact secondary derivatives of the armatured type. There is considerable inter-specific variation in hemipenial structure which is described systematically. In some cases this involves differences in size, asymmetry and simplification, which may arise as physical isolating mechanisms and is useful in distinguishing otherwise very similar species, particularly in the genus Mesalina (p. 1253). Other shared derived hemipenial features provide useful information about relationships between species and higher taxa and a summary of the hypotheses that they support is given (p. 1254).

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

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Estimates of phylogeny may allow historical events to be reconstructed even without a fossil record. The reliability of such interpretations depends not only on the robustness of the phylogeny but also on its topology. Changes in individual features can be traced and general histories of groups developed and compared with each other. Results are ofter surprising, for Instance the sophisticated tail shedding mechanism of lizards turns out to be a primitive feature that has been lost many times. Similarly, ecological analogues may have developed their common characaters in quite different orders. Phylogenies also provide a way of recognizing constraints and the effects of history on present ecological and behavioural patterns. When using anatomical characters the quality of the apparent phylogenies produced may be related to ecological history: expansion of a group along an ecological continuum into increasingly demanding niches ina small geographical area tends to produce a robust phylogeny (for instance in Meroles), while this is often not so for widely distributed groups that occupy a more modest range of niches (such as Pedioplanis and Podarcis). Non-morphological data may not show this tendency, but can have their own problems. Lacertids can be referred to clade with many successive branches in Africa and the Saharo-Eurasian arid zones and a less resolved probably paraphyletic complex of more primitive forms in the Mediterranean and wider Palaearctic areas. The African-Eurasian clade shows a general trend towards ground-dwelling and increasingly arid habitats but is ecologically variied. These animals are important to the study of Mediterranean forms because they provide eco-morphological parallels to them (for instance to Algyroides, Psammodromus and the archaeolacertas) and help form a basis for testing hypotheses about function. Although it is possible to recognise a number of distinct clades among the Mediterranean and Eurasian forms, relationships within and between these are often much less well substantiated. If results are available in time, current work on mitochondrial DNA sequencing will be discussed.

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

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

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We carried out a morphometric study of the skull throughout postnatal ontogeny in 14 species of Iberian lacertid lizards belonging to the genera Acanthodactylus, Algyroides, Psammodromus, Podarcis, Zootoca, Timon and Lacerta s. latu with the aim of elaborating a key of identification for the studies species. A total of sixty one characters showing interspecific variation are identified. The characters are defined in articulated and disarticulated skulls.

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Intra- und interspecific variation are analysed fpr the post-natal skull of the lacertid lizards Gallotia galloti, Actnhodactylus erythrurus, Algyroides marchi, Lacerta monticola, Lacerta vivipara, Podarcis bocagei, Podarcis hispanica, Podarcis muralis, Psammodromus algirus and Psammodromus hispanicus. Individual variations identified include: the prsence/absence, number and morphology of some structures, mainly in the lacrimal, sclerotic ossicles and macilla; the number and location of foramina and the degree of ossification of some processes in chondrocranial bones. No differences bones. No differences have been found between the sexes with respect to presence and morphology of the bones but some species show sexual dimorphism in the size and robustness of the head and this may be reflected in the individual. The most substantial variations seen are ontogenetic: the appearances of new traits; development of articulations; differences between species in the timing of the stages of development of a given structure; changes in the relative position of some cranial elements; and the degree of calcification or ossification of processes. A total of 63 characters showing interspecific variations are identified. Some have not previously been described while others used in previous studies are redefined here on the basis of new morphological information obtained.

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Males of many lizard species have longer tails than similarly-sized females. We hypothesized that this dimorphism is induced by a longer non-autotomous tail part in males, which is associated with the presence of the copulatory organs at the tail base, and presumably reduces the males` ability to escape predation by tail shedding. A compensatory mechanism would be an increase of total tail length in males, to achieve equal lengths of the autotomous tail part in both sexes. A critical prediction of this `morphological constraint` hypothesis is that the extent of dimorphism in total tail length increases with the magnitude of sexual differences in length of the non-autotomous tail base. We tested this prediction through a comparative study in a small clade of lacertid lizards. Within each of nine species, sexual differences in length of the non-autotomous tail base and in total tail length do not change with body size. All species, except one, exhibit a clear male-biased dimorphism in length of the non-breakable tail base. In all species studied, males have longer tails than females. We used the method of phylogenetically independent contrasts to explore the interspecific relation between dimorphism in length of the tail base and sexual differences in total tail length. Contrary to our prediction, we found no evidence for a positive correlation between the extent of dimorphism in both traits. Thus, constraints imposed by the male copulatory organs on tail autotomy do not seem to be a significant factor in the evolution of dimorphism in tail length in this clade of lacertid lizards.

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

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The list of amphibians and reptiles living in the Provence-Alpes-Côte d’Azur constitutes an inventory of up-to-date knowledge on herpetofauna.

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We describe a new location of Psammodromus edwardsianus in the Aragon southern area.

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We have studies the femoral pores of 387 subjects belonging to five species of Lacertidae family (Reptilia) of the región of Andalucia (Spain): Lacerta hispanica Steind., Lacerta lepida Daud., Psammodromus algirus L., Psammodromus hispanicus Fitz., Acanthodactylus erythrurus Schinz. We have observed that these femoral pores have a big importance in the sexual dimorphism which can be observed in the subjects after the birth. Possibly they take part in the phenomenous connected with the territorial behaviour of animals.

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Reproductive behaviour in Psammodromus hispanicus and Psammodromus algirus is pictu- red. P. hispanicus males grasp the dorsal skin of the females with their jaws during copula- tion of 40 s; males of P. algirus seize the females in the side of the neck and copulate for 8 min. Oviposition of both species is described. Dimensions of eggs, and duration of incu- bation and measurements of juvenile P. hispanicus are given. Specimens of the latter species are sexually mature within the year. In captivity they may live for three years or more. The change of juvenile colouration into adult pattern of P. hispanicus is reported. Rhythmical snout pushing as an expression of submissive behaviour; head shaking to ward off threats; and fights between males are described for P. hispanicus.

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The interspecific relationships among female size, clutch size, egg size, and hatchling size were examined for 64 European lacertids. The eggs of all species increased linearly in both linear dimensions and mass during incubation. Across species initial egg mass was positively correlated with juvenile mass, with an allometric relationship exponent of 0.87. Initial egg mass across species increased proportionally with female mass to the power of 0.57. Moreover, an increase in maternal mass was also accompanied by an increase in clutch size. The number of eggs per clutch across species scaled with female mass to the power of 0.39. Removing the effect of female mass resulted in a negative correlation between egg mass and clutch size. Species for which the average egg size was lower than expected on the basis of female mass, tended to have larger relative clutch size. The total egg mass per clutch was about one third of female mass (exponent 0.94).

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An observation of the Spanish Psammodromus in the Provence is described. The author visited the area frequently for eighteen years, but didn’t see this species before. July 2002 he saw a female of this species at the edge of a field of a grapery near Le Luc, close to shrubs (macchie). The lizard was very timid and couldn’t be observed afterwards. It is not clear whether this observation is related to an enlargement of the known range.

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Psammodromus algirus y Psammodromus edwardsianus (Reptilia: Lacertidae) son hospedadores en España de nematodos pertenecientes al género Parapharyngodon (Nematoda: Oxyuridomorpha, Pharyngodonidae). Una atenta observación de los rangos de distribución, tanto de parásitos como de hospedadores, sugiere que los primeros pudieron haber llegado accidentalmente a España desde el Nuevo Mundo en barcos de la “Carrera de Indias”.

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Habitat selection of Psammodromus algirus and P. hispanicus was studied along a successional vegetation gradient in Southern Spain. The two species select areas with a high cover of herbs and bushes at ground level and at a height of 10 cm above ground. Whereas P. algirus avoids sandy substrates and selects wooded areas, P. hispanicus, which does not enter the forest, occupies areas with a denser cover of low shrubs and herbs. Therefore, and despite their coincidence in the centre of the habitat gradient and hence their high overlap in habitat use, both species appear to be segregated, with P. hispanicus having a narrower spatial niche than P. algirus. The observed differences in habitat selection are discussed in the light of differences in morphology, reproductive constraints and predator avoidance. We conclude that interspecific competition plays a negligible role in the habitat segregation of the two species.

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Psammodromus hispanicus is a small lacertid lizard which ranges all the Iberian Peninsula, excepting the north and the Pyrenean Mountains (BARBADILLO 1987) and the SE of France (FRETEY 1987). It is nominally protected by Spanish law (Boletin Oficial del Estado 56:4999-5002,6.3.1981). Recent studies have analyzed its ecological trends and natural history (PASCUAL & PEREZ-MELLADO 1987, 1989, POLLO & PEREZ-MELLADO 1988,1989,1990, CARRETERO & LLORENTE, in press). It occurs mainly in the Mediterranean garigue or in more open areas, around patchy dense vegetation (ARNOLD 1987, CARRASCAL et al. 1989). It is common to see animals moving quickly from one plant (20-30 cm high) to another. In the NE of the Iberian Peninsula, the most dense populations are located in the coast, particularly in sandy areas. The development of the Herpetological Atlas of Catalonia (unpublished data) and Languedoc-Rousillon (GENiEZ & CHEYLAN 1987) have allowed us to prove that the range of distributionof this species is not continuous. At the present, French and Spanish populations are not connected (Fig. 1) but they were probably a continuumin the recent past (FELIXpcrs. com.). The northernmost Spanish population is located in Pals Beach (UTM 31TEG25) where P. hispanicus and its congeneric species P. algirus live together. The south- ernmost French records are close to the Spanish frontier. The intermediate zone (Alt Empordi region) has some favourable sites to be occupied, such as the coastal dunes between Muga and Fluvi^ rivers (Fig. 2). P. algirus is present there and has been previously studied (CARRETERO & LLORENTE 1989). The psammophilous vegetation (Theucrium, Echinophora, Ammophila, Pan- cratium} and the habitat structure are quite similar to other coastal locations of the Spanish Psammodromus. Indeed, a record near that site was published at the beginning of this century (MALUQUER 1916). The above facts have induced us to believe that the species has become extinct recently. Probably the tourist industry (destruction of psammophilous vegetation near beaches) is responsible. Now, this zone belongs to the Pare Natural dels Aiguamolls de 1`Empordil created in 1985 and it receives complete protection. Since the creation of the park, the possibility of rcintroducing the species has been planned with two consecutive objectives: 1.- Management: re-creation of a stable populationof Psammodromus hispani- cus. 2. - Ecology: employment of this reintrodnotion as a tool to know how both Psammodromus species use the same ecological resources. Only the first point is being carried out and it is reported here. The next step was to decide when and where to obtain the animals. The reproduction of the species has been previously studied in the NE of Spain (CARRETERO & LtORENTE in press). This lizard has a one-year cycle (Fig. 3). Young are born from July to September. They grow up very fast, becoming adults during the next spring. There is no winter diapause period. The life is very short. Only 5% of the adult lizards can breed in the second ye.ir. Two clutches have been detected in large females and they are reflected in the ovarian cycle (Fig. 4).

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The reproduction of a population of Psammodromus hispanicus was studied on a coastal sand area of NE Spain and compared with other populations. Sexual maturity is reached in the first spring, and there are no subadults. Reproductive activity occurs in March to July. The beginning of reproduction depends on the spring temperature increase but the end is produced by an endogenous rhythm. Eggs size, clutch size and length of the hatchlings are very close to those of other populations. The major difference is that a second clutch can be laid by the largest females. This is the first such record in the species.

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The variation in the reproductive cycle of three populations of the lizard Psammodromus algirus living in coastal sandy areas of NE Spain was analyzed. In contrast with those in continental climates, they showed evidence of reproductive stress: the reproductive season began sooner and was longer. Moreover, body size was smaller and sexual size dimorphism did not arise. Only the end of the breeding period was relatively constant. Nevertheless, these different reproductive traits seem to follow a common trend. In general, no important reproductive differences were found among the three coastal areas. Females laying two clutches have been detected in one locality. Clutch size was correlated with female size. The high phenological plasticity of this species is associated with its wide ecological niche.

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

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The Psammodromus hispanicus species group has been recently shown to include three lineages that differ in morphology (San-Jose et al. 2012), have largely parapatric range but exhibit little evidence of historical gene flow (Fitze et al. 2011), leading to the recognition of these three lineages as distinct species (Fitze et al. 2012). The eastern species can be unambiguously associated with the nomen Lacerta edwarsiana Dugès 1829, as the detailed information in Dugès (1829) leaves no doubt that he describes as Lacerta edwarsiana the local member of the P. hispanicus complex, and the type locality is the `bas Languedoc`, an area of France equivalent to the lowland parts of the current Languedoc region where the only member of the complex is the eastern lineage. The types of Psammodromus edwarsianus have not been traced as far as I am aware, but given the lack of uncertainty regarding allocation of this nomen to the eastern lineage of the P. hispanicus complex this has no nomenclatural consequence. Two nomenclatural issues remain in this species group however: the aim of this note is to solve them.

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There are many factors affecting population dynamics, both abiotic and biotic. One of the most important biotic factors affecting different ecological aspects of a species is competition. But there is relatively little research focussing on the effect of competition on population dynamics. Thus, we studied population dynamics of three insectivorous lizard species (Acanthodactylus erythrurus, Psammodromus algirus and Psammodromus edwardsianus) which cohabit in our study area. We carried out periodic censuses and samplings during 2015 and 2017. Our results seem to confirm that a high level of competition can alter some aspects of population dynamics. Also body size seems to play a role, as most adjustments we detected were related to small size classes and the most affected species was P. edwardsianus. Some possible responses to competition we detected were an advance of the reproduction period, increased growth rates and biasing the sex ratio notably towards males (as higher testosterone levels lead to more aggression and this improves success rates in interspecific encounters). However, our study area is highly anthropized and further research is needed to discern if these adjustments are really exclusively related to competition or if human disturbance also plays a role.

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Trophic ecology is one of the most important aspects of a species’ ecology and it is crucial when trying to assess how different species interact in an ecosystem. Showing how the diet intraspecifically changes in different body sizes, sexes or habitats and how trophic resources are shared interspecifically allows to understand how the ecosystem functions. With this objective, we studied the diet and trophic ecology of Acanthodactylus erythrurus, Psammodromus algirus and Psammodromus edwardsianus, which are in syntropy in the Albufera de Valencia Natural Park. We collected a total of 485 faecal samples and could identify 18 different prey groups. The main prey groups in A. erythrurus were Hymenoptera, Coleoptera and Hemiptera; in P. algirus, Lepidoptera, Arachnida, Coleoptera, Diptera, Hemiptera and Orthoptera; and in P. edwardsianus, Coleoptera, Hemiptera and Arachnida. Our results show that A. erythrurus seems to be the most specialized species, having the lowest values of trophic niche amplitude and the lowest intraspecific diet variation. Although this species is known to feed mainly on ants, our results indicate that mirmecophagy seems to be especially important in juveniles. Indeed we observed in all three species a niche separation between small and big individuals, indicating that diet changes with age. We could also evidence a resource partitioning between A. erythrurus and P. algirus, presenting a niche overlap of only 51.6%. In conclusion, there are various factors playing together shaping the diet of a species in a certain environment: behavioural, ecological, environmental and physiological factors.

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Three related and morphologically similar small Psammodromus species (P. hispanicus complex) occur in the Iberian Peninsula and Mediterranean France, Psammodromus hispanicus, Psammodromus occidentalis, and Psammodromus edwarsianus. Despite recent works, overall ranges of the species remain partially uncertain, in particular for Westernmost part of Iberia and South-western France. Here we include data from 15 additional specimens that were sequenced for part of the mitochon-drial gene Cytochrome b to allow comparison with published data. We provide genetic confirma-tion that P. edwarsianus is the species of the complex occurring in France, and that P. occidentalis occurs on coastal and inland Portugal. Within P. occidentalis notable diversity occurs for this marker, which warrants further investigation. Some areas, particularly southern Portugal, still need assessment to clarify the distribution of these mostly cryptic species across the region.

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A new species of lacertid lizard of the genus Psammodromus is described from the Iberian Peninsula. Genetic and recently published phenotypic data support the differentiation of Psammodromus hispanicus into three, and not as previously suggested two, distinct lineages. Age estimates, lineage allopatry, the lack of mitochondrial and nuclear haplotype sharing between lineages, ecological niche divergence, and the current biogeographic distribution, indicated that the three lineages correspond to three independent species. Here, we describe a new species, Psammodromus occidentalis sp. n., which is genetically different from the other sand racers and differentiated by the number of femoral pores, number of throat scales, snout shape, head ratio, green nuptial coloration, and number of supralabial scales below the subocular scale. We also propose to upgrade the two previously recognized subspecies, Psammodromus hispanicus hispanicus Fitzinger, 1826 from central Spain and Psammodromus hispanicus edwardsianus (Dugès, 1829) from eastern Spain, to the species level: Psammodromus hispanicus stat. nov. and Psammodromus edwardsianus stat. nov. Given that the holotype of Psammodromus hispanicus was lost, we designate a neotype. We also analysed museum specimens of P. blanci, P. microdactylus and P. algirus to describe differentiation of the Psammodromus hispanicus lineages/species from their closest relatives.

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Background Genetic, phenotypic and ecological divergence within a lineage is the result of past and ongoing evolutionary processes, which lead ultimately to diversification and speciation. Integrative analyses allow linking diversification to geological, climatic, and ecological events, and thus disentangling the relative importance of different evolutionary drivers in generating and maintaining current species richness. Results Here, we use phylogenetic, phenotypic, geographic, and environmental data to investigate diversification in the Spanish sand racer (Psammodromus hispanicus). Phylogenetic, molecular clock dating, and phenotypic analyses show that P. hispanicus consists of three lineages. One lineage from Western Spain diverged 8.3 (2.9-14.7) Mya from the ancestor of Psammodromus hispanicus edwardsianus and P. hispanicus hispanicus Central lineage. The latter diverged 4.8 (1.5-8.7) Mya. Molecular clock dating, together with population genetic analyses, indicate that the three lineages experienced northward range expansions from southern Iberian refugia during Pleistocene glacial periods. Ecological niche modelling shows that suitable habitat of the Western lineage and P. h. edwardsianus overlap over vast areas, but that a barrier may hinder dispersal and genetic mixing of populations of both lineages. P. h. hispanicus Central lineage inhabits an ecological niche that overlaps marginally with the other two lineages. Conclusions Our results provide evidence for divergence in allopatry and niche conservatism between the Western lineage and the ancestor of P. h. edwardsianus and P. h. hispanicus Central lineage, whereas they suggest that niche divergence is involved in the origin of the latter two lineages. Both processes were temporally separated and may be responsible for the here documented genetic and phenotypic diversity of P. hispanicus. The temporal pattern is in line with those proposed for other animal lineages. It suggests that geographic isolation and vicariance played an important role in the early diversification of the group, and that lineage diversification was further amplified through ecological divergence.

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We describe six new locations for Psammodromus edwardsianus (Dugès, 1829) in Catalonia, after campaign of research on the species.

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Karyotype data are presented for 12 species of lacertid lizards. Of these, 4 Acanthodactylus, 2 Eremias, 1 Ophisops, and 4 Lacerta have n = 19, with all chromosomes acrocentric. This is the typical lacertid karyotype, as reported in previous literature. One species, Lacerta parva, is quite different from all the others. The haploid number is 12, consisting of 7 pairs of metacentric macrochromosomes and 5 pairs of microchromosomes. The fundamental number (number of arms in the karyotype) is the same in L. parva as in all the other species studied. The possibility of female sex chromosomal heteromorphism is raised, but data are not sufficient to confirm this.

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In recent years there has been an exponential increase in the amount of published DNA sequences, as it has become one of the most popular sources of data for reconstructing phylogenies. A number of studies have attempted to resolve the relationships of groups of lacertids using mitochondrial DNA sequence data, in particular Podarcis and Gallotia (e.g. THORPE et al., 1994; 1996; GONZALEZ et al., 1996; FU et al., 1997; HARRIS, 1997; HARRIS et al., 1998a; 1999; CASTILLA et al., 1999a,b). Reassessment of these data suggest some alternative phylogenies to those previously published, and also highlight possible problems. Two studies have attempted to unravel the phylogeny of the whole family using mitochondrial DNA sequences (FU, 1998; HARRIS et al., 1998b). By combining these essentially congruent data sets, and reanalysing levels of support for the phylogenies produced, a more complete estimate of relationships across the family is produced. This is then compared to alternative sources of evidence, primarily morphological (ARNOLD, 1989), biochemical (MAYER & BENYR, 1994) and karyological (OLMO et al., 1993). Reasons for discrepencies are discussed, as are the relative advantages and disadvantages of the different sources of phylogenetic inference.

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

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

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The knowledge of a species’ population structure is essential for the development of adequate conservation actions as well as for the understanding of its evolution. The population structure is unknown in all species of the Genus Psammodromus, including the Western Sand Racer (Psammodromus occidentalis; a recently described species), the Edward’s Sand Racer (P. edwardsianus) and the Spanish Sand Racer (P. hispanicus). In this article, the genetic variability and population structure of Psammodromus edwardsianus, P. hispanicus, andP. occidentalis were studied in the Iberian Peninsula covering their natural geographic distribution. Mitochondrial DNA showed genetically different units in all species with higher genetic variability in their southern populations (latitudinal variation). Genetic differentiation was different among species and contrasted to those of species with similarcharacteristics. Our results therefore highlight the importance of species-specificstudies analysing population structure.

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While ecological effects on short-term population dynamics are well understood, their effects over millennia are difficult to demonstrate and convincing evidence is scant. Using coalescent methods, we analysed past population dynamics of three lizard species (Psammodromus hispanicus, P. edwardsianus, P. occidentalis) and linked the results with climate change data covering the same temporal horizon (120 000 years). An increase in population size over time was observed in two species, and in P. occidentalis, no change was observed. Temporal changes in temperature seasonality and the maximum temperature of the warmest month were congruent with changes in population dynamics observed for the three species and both variables affected population density, either directly or indirectly (via a life-history trait). These results constitute the first solid link between ecological change and long-term population dynamics. The results moreover suggest that ecological change leaves genetic signatures that can be retrospectively traced, providing evidence that ecological change is a crucial driver of genetic diversity and speciation.

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Many lizards produce chemical secretions that may be used as pheromones in reproductive behavior, but only a few studies have identified chemical compounds in secretions. By using GC–MS, we found only 20 lipophilic compounds in femoral glands secretion of male lizards, Psammodromus hispanicus. Main compounds were six steroids (mainly cholesterol and campesterol) and seven n-C9 to n-C18 carboxylic acids (especially dodecanoic acid), and minor components were six alcohols between C16 and C29 and squalene. We compared these chemicals with those previously found in secretions of the closely related sister species Psammodromus algirus and other lizard species.

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

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

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This note describes general patterns of winter activity in lizard populations along a latitudinal gradient with an approximate range of twelve degrees (26 to 38°N) in southern Europe and North Africa. We make use of qualitative data gathered through the last ten years in the area, as well as much more accurate quantitative data obtained from periodic censuses covering the whole year cicle in certain populations. Results allow us to impute a great importance to the activity observed in the unfavourable season. This activity seems to be an attribute at the population level and shows a great variability between and within species and populations.

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Aim To infer the biogeographical and evolutionary history of the Western Mediterranean Psammodromus lizards with the aim of assessing the role of vicariance and dispersal on the cladogenetic events within the palaeogeological dynamics of the Strait of Gibraltar. Location North Africa and Western Europe. Methods We built a dataset including all six species of Psammodromus using mitochondrial (12S, cytb, nd4) and nuclear (acm4, mc1r, pomc) gene fragments. Species tree and concatenation methods were used to infer phylogenetic relationships and divergence times. Phylogenies were used for biogeographical inference using S-DIVA, DEC and BBM. Results Psammodromus probably originated in Iberia, with P. algirus diverging early. The ancestor of the African P. blanci and P. microdactylus dispersed to Africa through the Betic-Rif massif, c. 10 Ma. The cladogenetic events within Africa and Iberia were probably due to vicariance mediated by habitat and climatic changes at the end of the Miocene (P. blanci and P. microdactylus) and during the Pliocene (P. occidentalis, P. hispanicus and P. edwardsianus). Psammodromus algirus shows three lineages, two in Iberia and one in Africa, the latter originated following a transmarine dispersal during the Middle Pleistocene (1.5 Ma). Main conclusions Over-sea dispersal has played a major role in intercontinental exchange and divergence in Psammodromus, with two dispersal events towards Africa that occurred 10 and 1.5 Ma resulting in the African lineages. This study, combined with previous literature, provides compelling evidence that major biotic exchanges took place across the Strait of Gibraltar well before or long after the land connection during the Messinian Salinity Crisis (MSC, 5.9–5.33 Ma). These findings suggest caution in the application of the relatively short event of Atlantic flooding at the end of the MSC as cause for divergence in molecular clock calibrations, which is a popular approach in literature.

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The genus Psammodromus (Lacertidae) comprises six species. Three of the species are morphologically similar, phylogenetically closely related, and are distributed in the Iberian Peninsula (besides P. algirus): P. occidentalis (Western Iberia), P. hispanicus (Central Spain), and P. edwarsianus (Eastern Spain). Previous studies have shown these three species to have allopatric distributions in Iberia, though there are still many areas such as Andalucia where it is unclear which species of Psammodromus occurs. In this study we present nine new records of Psammodromus from key places in southern Spain. Since the three Iberian species are difficult to differentiate by morphology, specimens were genetically identified using mitochondrial DNA data and compared to all six species of the genus. Here we present an updated distribution map of Psammodromus specimens identified with molecular methods, increasing the ranges of all three species. We also propose wider regions of potential sympatry between P. occidentalis and P. hispanicus, and P. edwarsianus and P. hispanicus than previously recorded.

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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 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 .

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Male and female Psammodromus hispanicus from southern Europe were acclimated to four seasonal conditions of photoperiod and night time temperature. During the dark period, the lizards` body temperatures fell to ambient air temperature but during the light period the lizards were allowed to thermoregulate behaviourally and at such times the lizards` mean body temperature varied from 29.0°C to 32.6°C. The resting metabolic rate of these lizards was measured in 5°C steps from 5°C to 30°C or 35°C. Sexual condition had little effect on resting metabolic rate, but at low temperatures lizards acclimated to winter or spring seasonal conditions had lower resting metabolic rates than those acclimated to summer or autumn conditions. At temperatures above 20°C seasonal acclimation had no effect on resting metabolic rate. It is considered that the reduction in low temperature metabolic rate in spring and winter is induced by low night time temperatures and serves to conserve energy during those seasons when lizards must spend long periods at low temperature without being able to feed.

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Lacertids show three types of conspicuous lateral colour patches (i.e. ultraviolet-blue, ultraviolet- green, and ultraviolet-yellow), that appear repeatedly as alternative phenotypes across the phylogeny of this lizard family. Theory predicts that chromatic signals have evolved to be more conspicuous in their own visual ecosystem. Thus, there should be a relationship between a species’ habitat and the type of conspicuous coloration it shows, considering all observers that might exploit these conspicuous colour patches (i.e. conspecifics, predators). In this work we tested if three sympatric species, Psammodromus algirus, P. edwardsianus, and Acanthodactylus erythrurus (which show the three alternative phenotypes), (1) present differences in conspicuousness, and (2) are segregated by habitat (i.e. light spectral quality, openness, and available refuges). We hypothesized a relationship between the degree of conspicuousness and microhabitat selection, with the most conspicuous species occupying the most vegetated and refuge-rich habitats, and the less conspicuous species occupying more open habitats with less refuges. Our results confirmed our initial hypothesis, and suggest that lizards with the most conspicuous colorations select those microhabitats in which they are less vulnerable to visual predators. These results agree with the idea of a direct relationship between habitat and colour signal design.

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

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We report on the reproductive cycle and reproductive characteristics of three oviparous lizard species: Psammodromus algirus, Psammodromus hispanicus and Acanthodactylus erythrurus from a sandy coastal area in SW-Spain. Maximum testicle activity in P. algirus is research between March-May. Oviposition occurs along the activity cycle until September, clutch size average 4.8 eggs (2-10) and larger females tend to produce larger clutches. In P. hoispanicus testicle reaches maximal volume during February-May. Oviposition occurs throughout April-September, and two clutches, averaging 3.6 egg (2-5), are produced annually. In A. erythrurus mean testicle colume reaches its maximum in March, oviposition occurs in two cycles: spring and summer, clutch size averages 4.4 eggs (3-6) and increases with female snout-vent length.

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Studies on thermoregulation and temporal distribution of Psammodromus algirus and Psammodromus hispanicus have been carried out in a coastal area of SW-Spain (Huelva province). The annual activity cycle of Psammodromus algirus is continue: from January to December; maximum activity level of adults and subadults is reached in May (air temperature between 25-30°C) and minimum during summer and winter months. Daily cycle displays two annual evolution models in relation to air temperature: 1) Unimodal pattern, with maximum during the midday (October and April); and 2) Bimodal pattern (display in U) with depressed activity during the warmer summer months at the midday (May-September). Such seasonal shifts in time of activity facilitate thermoregulation, body temperature (BT) of active Psammodromus algirus during the annual cycle averages 30.9°C (19.0-40.1, SD= 5.66, N= 188); relations between BT, air (AT) and substratum (ST) Temperature are significant (p<0.01): r= 0.6325 and r= 0.8262, respectively. Psammodromus hispanicus decrease its activity rhythm during nwinter; maximum activity level is reached in May, daily activity pattern is associated with air temperatures lower than 26.0°C, over these temperatures the number of observations decrease until reach the minimum at 17.00 (Mean European Time). Body temperature of active Psammodromus hispanicus during the annual cycle averages 32.6°C (28.8-36.6, SD= 2.34, N= 63); relations between BT, AT and ST are significant (p<0.01): r= 0.6056 and r= 0.4748, respectively.

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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.

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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.

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Home range and spatiotemporal activity are often lacking for small vertebrates that are difficult to mark individually and to monitor over sufficiently long-time scales to collect reliable information. This is particularly the case for the Edward’s Sand Racer Psammodromus edwarsianus, a small Mediterranean lizard that is almost threatened with extinction in France. In order to fill these gaps, we conducted a four-year mark-recapture survey (2019-2022) carried out on a 0.5ha quadrat composed of two distinct habitats of equivalent surface area (open rocky area and Aleppo pine forest). We estimated the 95% and 50% (core area) spring home ranges for 10 adult individuals (5 males and 5 females) using the Autocorrelated Kernel Density Estimation method. We assessed daily activity patterns of P. edwardsianus and tested whether these differed between the two habitat types using a Generalized Linear Mixed Model. The surface of spring home range varied greatly between individuals (for females, 95% AKDE range from 733m2 to 3340m2 and 50% AKDE range from 158m2 to 546m2. For males, 95% AKDE range from 4556m2to 7434m2and 50% AKDE range from 779m2 to 1658m2), and reached up to 290 times the value formerly reported in the literature with significantly larger spring home ranges for males than females. The activity of P. edwardsianus did not vary significantly between the two habitats, and regardless of habitat type, activity was highest in the morning and late afternoon. The data collected have enabled us to gain a better understanding of the displacement potential and the spatiotemporal activity patterns of P. edwardsianus. This study also provide methodological elements and advice for optimizing the monitoring of this species.

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Parasitic helminths are an almost universal feature of vertebrate animals, but reptiles are among the hosts with the mostdepauperate parasite communities. Biological traits of reptiles are considered to be among the key reasons that explain low helminthdiversity; therefore, insights from a wide range of reptile hosts are helpful to understanding the ecology of parasitic helminths. Weanalyzed helminth fauna in two lacertids,Psammodromus algirusandPsammodromus edwarsianus(Squamata: Lacertidae), and oneskink,Chalcides bedriagai(Squamata: Scincidae), three common species of Mediterranean woodlands that differ in their ecologicalconditions and in lifestyles that are linked to habitat use. We examined a total of 102P. algirus,27P. edwarsianus, and 23C. bedriagaifrom mountain landscapes in eastern Iberia. We found three helminth species, the nematodesParapharyngodon echinatusandSpauligodonsp., and the cestodeMesocestoidessp. We report new reptile hosts for two helminth species: the skinkC. bedriagaifor thenematodeP. echinatusand the lizardPsammodromus algirusfor the nematodeSpauligodonsp. We also provide the second record of thelarval forms of the cestodeMesocestoidessp. in the lizardP. edwarsianusfrom the Iberian Peninsula. Interestingly, prevalence ofinfection was much higher in the skink than it was in the two lacertid lizards. Therefore, a subterranean skink lifestyle may determinethe incidence of helminth parasites when compared to the ground-dwelling lizard species. Similar to other reptile hosts, the helminthfauna of our focal lizard species was poor and mainly composed by Pharyngodonidae nematodes that are often detected in insectivorousreptiles.

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The Spanish sand racer (Psammodromus hispanicus) has been recently split into three distinct species: P. hispanicus, P. edwardsianus, and P. occidentalis. Some morphological differences have been reported but there is as yet no description allowing unambiguous identification of the three species. Here, we describe differentiation in body measurements, scalation traits, and colour traits as well as in the degree of sexual dimorphism. Our results show that P. edwardsianus can be easily distinguished by the presence of a supralabial scale below the subocular scale, which is absent in the other two species. Psammodromus hispanicus and P. occidentalis can be distinguished by the number of femoral pores, throat scales and ocelli, and the relative width of the anal scale. The degree of sexual size dimorphism and sexual colour dimorphism substantially differs among species, suggesting that different scenarios of sexual and natural selection may exist for each species. Moreover, sexually selected traits (nuptial colouration, ocelli, and femoral pores) significantly differ among species, suggesting that visual and chemical communication may also differ among species. Such differences could prevent reproduction and gene flow at secondary contact zones, potentially reinforcing isolation and speciation within this group of lizards.

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Extreme climate events, together with anthropogenic land use changes, have led to the rise of megafires (i.e., fires at the top of the frequency size distribution) in many world regions. Megafires imply that the centre of the burned area is far from the unburnt; thus, recolonization may be critical for species with low dispersal abilities such as reptiles. We aimed to evaluate the effect of megafires on a reptile community, exploring to what extent reptile responses are spatially shaped by the distance to the unburned area. We examined the short-term spatiotemporal response of a Mediterranean reptile community after two megafires (>20.000 hectares) occurred in summer 2012 at eastern Spain. Reptiles were sampled during four years after the fire in burnt plots located at different distances from the fire perimeter (edge, middle, and centre), and in adjacent unburnt plots. Reptile responses were modelled with fire history, as well as climate and remotely sensed environmental variables. In total we recorded 522 reptiles from 12 species (11 species in the burnt plots and 9 in the unburnt plots). Reptile abundance decreased in burnt compared to unburnt plots. The community composition and species richness did not vary either spatially (unburnt and burnt plots) or temporally (along the four years). The persistence of reptiles in the burnt area supports their resilience to megafires. The most common lizard species was Psammodromus algirus; both adults and juveniles were found in all unburnt and burnt plots. This species showed lower abundances in burnt areas compared to the unburnt and a slow short-term abundance recovery. The lizard P. edwarsianus was much less abundant and showed a tendency to increase its abundance at burnt plots compared to unburnt plots. Within the megafire area, P. algirus and P. edwarsianus abundances correlated with the thermal-moisture environment and vegetation recovery regardless of the distance from the fire edge. These results indicate the absence of a short-term reptile recolonization from the unburnt zone, demonstrating that reptiles are resilient (in-situ persistence) to megafires when environmental conditions are favourable.

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

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

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

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We assess associations between chemosensory capacity and foraging mode within lacertid lizards. Species of Lacertidae differ considerably in indices of foraging mode, and therefore seem well suited to test the adaptive nature of a coupling between foraging and sensory ecology. We observed tongue-flick (TF) rates of members from eight species in the field and in experimental conditions with no prey stimuli, with chemical stimuli, with visual stimuli and with both chemical and visual stimuli. All species increased TF rates in response to both purely visual and purely chemical prey cues, and the increase was most pronounced when both types of stimuli were offered simultaneously. Absolute TF rates in experimental conditions differed considerably among species, and species means obtained in experimental situations correlated closely with those observed in the field. Species that spend a relatively large amount of their time budget actively searching for food tend to use their vomeronasal system more frequently. Although all species in our study retained the capacity of recognizing prey chemicals, our data corroborate the idea of a functional link between an active foraging style and the usage of chemical information.

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

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

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