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We found significant variations in the pattern of sexual dimorphism in body size and shape among and within three lacertid species (Podarcis muralis, P. melisellensis and Archaeolacerta oxycephala), which live within a very restricted area (Lake Skadar, Montenegro) and under similar ecological settings. Each species has a specific pattern of size and shape relations between the sexes that proportionally changes with increase in size of the males, contrary to the expectations of Rensch`s rule. Our data do not confirm unequivocally that the level of size differences between the sexes (SSD) is affected by the body shape of two different morphotypes (cylindric vs. flattened body), or by the insularity conditions (island size, distance from mainland and the presence of syntopic lizard species). However, SSD appears to be positively correlated with clutch size at the interspecies level of comparisons.

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We estimated changes in the components of phenotypic variance and covariance among seven metric traits over the first 90 days of growth in the sharp-snouted rock lizard (Lacerta oxycephala). The broad-sense heritability estimated from the data representing mostly size components of the seven morphometic traits over four age periods was about 90%. The average broad-sense heritability for the shape components of the same traits and age periods amounted to about 42%. By using bivariate and multivariate statistical methods, ontogenetic allometry coefficients have been estimated. The average heritability estimated for the bivariate allometric coefficients was about 69%. Genetic correlation structures were highly integrated within each age period and this integration increased from newborn lizards to lizards aged up to 90 days. For absolute measurements, phenotypic correlation structures were less integrated than genetic correlation structures, while for size-free measurements the opposite trend was observed.

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

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

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

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

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

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

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

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

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1. The signals that animals use to communicate often differ considerably among species. Part of this variation in signal design may derive from differential natural selection on signal efficacy: the ability of the signal to travel efficiently through the environment and attract the receiver`s attention. For the visual and acoustic modalities, the effect of the physical environment on signal efficacy is a well-studied selective force. Still, very little is known on its impact on chemical signals. 2. Here, we took a broad, phylogenetic comparative approach to test for a relationship between animals` signal chemistry and properties of their natural environment. Our study focused on lizards from the Lacertidae family. 3. We sampled 64 species across three continents and determined the lipophilic composition of their glandular signalling secretions using gas chromatography-mass spectrometry. For each species, an array of environmental variables of high temporal and spatial resolution was obtained from climate databases. 4. Species varied considerably in the overall richness (number of constituents) of their secretions, as well as in the relative contribution of the major chemical compound classes. Signal richness and the relative contribution of the respective compounds exhibited little evidence of phylogenetic relatedness, suggesting that chemical signals may change very rapidly. Neither insularity nor substrate use affected chemical signal composition, however, we found a strong statistical relationship between the chemistry of the lizards` secretions and aspects of the thermal and hydric environment they inhabit. 5. Species from `xeric` milieus contained high proportions of stable fatty acid esters and high molecular weight alcohols in their glandular secretions, which likely increase the persistence of secretion scent-marks. In contrast, species inhabiting `mesic` environments produced secretions of a high chemical richness comprising high levels of aldehydes and low molecular weight alcohols. This chemical mix probably creates a volatile-rich signal that can be used for long-distance airborne communication. 6. We argue that the observed variation in signal design results from differential natural selection, optimizing signal efficacy under contrasting environmental conditions.

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

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We studied impacts of the introduced small Indian mongoose Herpestes auropunctatus on the herpetofauna on six islands in the Adriatic Sea, Croatia, comparing abundances of reptiles and amphibians on three islands with the mongoose to those on three islands without the mongoose. We used four types of sampling surveys: distance-constrained surveys, visual encounter surveys, special searches and accidental trapping. The horned viper Vipera ammodytes and Balkan green lizard Lacerta trilineata were absent from two mongoose-infested islands (Korčula and Mljet) and rare on the third (Hvar); they were common only on the mongoose-free island where they had historically been present (Brač). The European green toad was absent from one mongoose-infested island, where it had historically been present and rare on the other two. It was common on two of the three mongoose-free islands. Other herpetofaunal species were either very scarce or completely absent on the three mongoose-infested islands. Most of these species also occur on the mainland but are already scarce there; some are strictly protected under Appendix II of the Berne Convention. The recent spread of the mongoose to the European mainland suggests the need for urgent control to protect vulnerable herpetofauna.

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n response to two previously published accounts, it is pointed out that successful incubation of lizard eggs does not primarily depend on the medium on or in which the eggs rest, nor on the incubator. Successful incubation is solely based on the two physi- cal variables of humidity (measured as water potential in kPa) and temperature. When these variables are within the proper range (-400/-150 kPa and 25-30°C) then the chief criteria are met. The inaccurate suggestion that the sex of lacertids is re- lated to incubation temperature is rejected because the gender of the young in this group of reptiles is determined genetically. The single report cited in the scientific literature that discussed TSD in a lacertid (Podarcis pityusensis) is refuted by consid- ering larger samples of incubated eggs. Similarly, reports that water potential of the substrate affects sex determination have also been discredited in the scientific lit- erature. The sometimes in hobby literature propa- gated short, artificially induced additional hibernation soon after the summer to bring on an additional reproductive period with egg production in November/December, is objectionable. The stress on the animals is such that the health of the lizards is com- promised. Moreover, among serious reptile enthusiasts there is only a limited demand for lacertids, discounting the need for so many offspring.

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

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During a visit to Osor on the island Cres, Croatia, in mid August 2016 I photographed one adult Dalmatolacerta oxycephala eating a stink bug of the (family Pentatomidae). Judging from the pertinent literature it is concluded that D. oxycephala eats a wide variety of invertebrates, including scorpions, beetles and possibly even centipedes. Ripe fruit is also eaten. D. oxycephala is comfortable with humans at Osor and in other habitats visited frequently by people, whereas it proves very shy in other habitats. At Osor one Hierophis gemonensis was apparently hunting D. oxycephala in a wall.

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

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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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Two previously known northernmost localities of the Sharp-snouted rock lizard in Montenegro were the entrance of the Komarnica Canyon (Nevidio), and the middle part of the Tara River Canyon (village Tepca). It was presumed that Komarnica and Piva canyons were once corridors for the spreading of this species from the Eastern Adriatic sub-Mediterranean area to the Tara River Canyon in the north. However, it had not been hitherto known if there were any other relict populations in the area. In the autumn of 2019 a localized population of D. oxycephala was discovered in the middle part of the Komarnica Canyon.

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The basin of Lake Skadar with its drainage area represents one of the Balkan Peninsula’s hotspots regarding amphibian and reptile fauna. The value of the batracho- and herpetofauna of the Lake Skadar region is reflected in a rich and diverse composition of species. This is undoubtedly the result of the basin’s specific geographic position and complex geological history. Recent studies revealed that Lake Skadar’s watershed is inhabited by 15 amphibian and 36 reptile species. Among these, one amphibian (Pelophylax shqipericus – EN) and two reptile species (Dinarolacerta mosorensis and Vipera ursinii, both VU) are considered to be globally threatened according to IUCN criteria. An additional three reptile species (Emys orbicularis, Testudo hermanni, and Elaphe quatuorlineata) could become threatened in the future. As an attractive tourist area, a significant part of the Lake Skadar region could be impacted by fast and intense anthropogenic changes. Therefore, preserving the network of suitable habitats, maintaining continuous monitoring, and investing in additional research are essential for maintaining this rich local amphibian and reptile diversity.

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Während eines einwöchigen Aufenthaltes im September 2006 auf der mitteldalmatinischen Insel Brač konnten drei Lacertidenarten nachgewiesen werden. Aufgrund der zu diesem Zeitpunkt recht hohen Tempersaturen fielen die Beobachtungen allerdings recht spärlich aus.

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

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We provide new and supplementary data on the distribution and natural history of reptiles on the island of Cres: Dalmatolacerta oxycephala (range extension of an alien introduced population); Emys orbicularis hellenica (status of the population in the southwestern parts of Cres); Hemidactylus turcicus (first record after 38 years); Trachemys scripta elegans (first record on Cres).

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Lizards belonging to the species Lacerta sicula, L. oxycerihala, and L. mellisellensis from numerous Adriatic Islands and from adjacent mainland populations show no variation in karyotype. All are characterized by the diploid number 2n - 3S consisting of 36 acrocentric macrochromosomes and 2 acroce trie microchromosomes.

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Twenty populations of Lacerta sicula, L. melisellensis, and L. oxycephala from the coast and islands of the Adriatic were studied electrophoretically. Genetic similarity measured by the Rogers coefficient (Rogers, 1972) ranges from 0.84-1.0 among populations of melisellensis, and from 0.88-0.99 among populations of sicula. Interspecific values range from 0.61-0.74 in this pair, and both have values of about 0.43 compared to oxycephala. There is circumstantial evidence for hybridization and introgression in one population. Among Lacerta, amounts of genetic variation are greater in (1) mainland populations than in putatively relict island populations, and (2) on large islands than on small islands. Some fringing islands that are separated by a short linear distance and shallow channel depth from the parent island show a `small island effect,` i.e., a precipitous drop in variation compared with the mother island but only if these fringing islands are tiny ($acksimeq leqslant$ 0.01 sq. km). Genetic drift is implicated. Larger fringing islands do not show an area effect and are genetically virtually identical to the parent island. An analysis of allele frequencies in populations on different islands suggests that most of the polymorphisms are being selected, but that some (e.g., esterases) have a pattern consistent with the neutraldrift hypothesis. Stepwise multiple regression was used to analyze the geographic correlates of mean heterozygosity. The results, like those of a previous study of island lizards, are consistent with the predictions of the time-divergence theory of variation, but do not allow firm conclusions. The theory predicts that the smaller (more ecologically distinct) and the older an island, the lower the heterozygosity in its population. It is argued that directional selection is the main force eroding genetic variation; the smaller an island, the more directional selection will prevail over stabilizing selection; the older an island, the longer such selection has operated. Very small islands, where genetic drift appears to reduce variability, were excluded from most of the regression studies.

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Thermal and trophic niche relationship between podarcis melisellensis and Lacerta oxycephala were studied from island of Vis in June and September 1997. We recorded 289 cloacal temperatures and related substrate and aur temperatures considering sex and age categories. It seems that p. melisellensis posses a certain independence from microhabitat temperatures as in many reptiles adaptet to life in desert regions. Females and subadults of L. oxycephala showed similar patterns. A study of diet showed that the most important prey groups for both species were Formicoidea. Food niche overlap between species was very high.

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Thermal and trophic niche relationship between podarcis melisellensis and Lacerta oxycephala were studied from island of Vis in June and September 1997. We recorded 289 cloacal temperatures and related substrate and aur temperatures considering sex and age categories. It seems that p. melisellensis posses a certain independence from microhabitat temperatures as in many reptiles adaptet to life in desert regions. Females and subadults of L. oxycephala showed similar patterns. A study of diet showed that the most important prey groups for both species were Formicoidea. Food niche overlap between species was very high.

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Diversity of amphibians and reptiles, as well as threat factors were investigated during last decade in the wide area of the Vrmac hill. Field methodology included active searching and visual observation of species along defined transects. In total, 29 species were recorded, what comprises about 54% of amphibian and reptile fauna known for Montenegro. Main threat factors for amphibians and reptiles in the area of Vrmac hill are: urbanization, habitat fragmentation and destruction, fires, illegal landfills, waste waters, direct killing by locals and road killing.

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Diversity of reptile fauna, as well as threat factors were investigated during April and May 2019 in the settlement Mareza. Field methodology included active searching and visual observation of species along defined transects. In total, 20 reptile species were recorded, what comprises about 55 % of reptile fauna known for Montenegro. According to obtained results, area of Mareza can be considered as important reptile diversity hotspot in Montenegro. First finding of Xerotyphlops vermicularis in this area contribute to knowledge of its distribution range. Three identified species (Emys orbicularis, Testudo hermanni and Elaphe quatuorli neata) are listed on Annex II of the Habitat Directive. None of identified species are vulnerable, endangered or critically endangered, upon the IUCN Red List. Almost all identified species are protected on national level. Main threat factors for reptile fauna in the area of Mareza are: urbanization, habitat fragmentation and destruction, fires, illegal landfills, direct killing by locals and road killing.

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This paper presents an updated checklist of Amphibians and Reptiles of Montenegro. The checklist contains 52 species (including two species complexes), compiled from published literature and personal field surveys. Species complexes are the Anguis fragilis complex and the Lacerta viridis complex. Trachemys scripta is introduced, while Podarcis siculus is potentially introduced species. Migratory species include three sea turtles (Caretta caretta, Chelonia mydas and Dermochelys coriacea). Additionally, there are four potentially present species (Proteus anguinus, Mediodactylus kotschyi, Tarentola mauritanica and Podarcis tauricus), which presence is doubtful or still not confirmed with certainty in Montenegro. National and international protection as well as conservation status of Amphibians and Reptiles is given.

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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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A remarkable hindleg regeneration was observed in a Sharp-snouted Rock Lizard (La- certa oxycephala). The hindleg was replaced by a scaled elongate and conical appendage. The moving lizard kept the regenerate motionless. The specimen apparently was able to catch enough food in nature. Comparable regenerations were found in Caucasian Rock Lizards, in two L. saxicola brauneri and in one L. raddei raddei. Always a hindleg was rege- nerated.

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In 1981/2 during five journeys to Yugoslavia distributional, ecological and systematic data were collected for the following species, mainly in the coastal area: Salamandra salamandra, Triturus cristatus, T. vulgaris, Bombina variegata, Bufo bufo , B. viridis, Hyla arborea, Rana dalmatina, R. latastei, R. ridibunda, Emys orbicularis , Testudo hermanni, Anguis fragilis, Ophisaurus apodus, Lacerta oxycephala , L. trilineata, L. viridis, Podarcis melisellensis, P. muralis , P. sicula, Coluber viridiflavus, Malpolon monspessulanum, Natrix natrix .

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

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

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

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Males of a Croatian population of the lacertid lizard Podarcis melisellensis exhibit a striking polymorphism, with coloration of the throat and abdomen ranging from completely white, to yellow or orange. In a first attempt to explore the potential ecological and evolutionary significance of this polymorphism, we compared the three forms of males in aspects of their morphology, whole-animal performance, behavior, and ecology. Orange males are, on average, larger in snout-vent length and have disproportionately larger heads than either white or yellow males. This is reflected in orange males having higher bite force capacity and theoretically an increased access to harder prey. Residual limb length, maximal sprint speed and maximal exertion do not differ among color morphs. Body temperatures in the field are similar in the three morphs, but yellow males are caught at sites with slightly higher air temperatures than are orange and white males. Behavioral observations show no differences in time budgets or in the timing of activities among morphs. Microhabitat use is also similar in the three color morphs, but orange males were more often initially seen on rocky substrates. Our findings suggest that the observed polymorphism likely does not originate from a divergence in niche or use of resources, but possibly reflects an underlying polymorphism in mating tactics.

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We present the distribution of amphibians and reptiles in the Bjelopavlici region (central Montenegro) based on our field work records and of literature data. Twelve species of amphibians and 25 species of reptiles were recorded at 49 localities of which 57 % records has not been previously published. The greatest diversity of batracho- and herpetofauna was recorded at sites in the river Zeta plain. Our study provides the important information for assessments of diversity of batracho- and herpetofauna of Montenegro and their conservation.

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

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A key assumption in evolutionary studies of locomotor adaptation is that standard laboratory measures of performance accurately reflect what animals do under natural circumstances. One widely examined measure of performance is maximum sprint speed, which is believed to be important for eluding predators, capturing prey, and defending territories. Previous studies linking maximum sprint speed to fitness have focused on laboratory measurements, and we suggest that such analyses may be appropriate for some species and intraspecific classes, but not others. We provide evidence for a general inverse relationship between maximum laboratory sprint speed and the percentage of maximum capacity that animals use when escaping from a threat in the field (the model of locomotor compensation). Further, absolute values of field escape speed and maximum laboratory speed are not significantly related when comparing across a diverse group of Anolis and lacertid lizards. We show that this pattern of locomotor compensation holds both within (i.e., among intraspecific classes) and among lizard species (with some exceptions). We propose a simple method of plotting field escape speed (y-axis) versus maximum laboratory speed (x-axis) among species and/or intraspecific classes that allows researchers to determine whether their study organisms are good candidates for relating laboratory performance to fitness. We suggest that species that reside directly on, or near the `best fitness line` (field escape speed = maximum laboratory speed) are most likely to bear fruit for such studies.

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Bosnia and Herzegovina plays an important role for the diversity of herpetofauna of the Balkans. The composition of the amphibian and reptile fauna has been influenced by both continental and Mediterranean climate. So far, 18 species of amphibians and 29 species of reptiles have been found to inhabit the territory, but due to the overall neglected faunistic research of this region in the last decades, this is presumably not the final number. At 31 localities we observed 12 species of amphibians and 17 species of reptiles representing 66.7 and 58.6% of the total known diversity of amphibians and reptiles, respectively. We recorded new distribution data for 8 species of amphibians and 9 species of reptiles. A zoogeographic analysis showed that the herpetofauna of Bosnia and Herzegovina could be classified into at least 12 chorotypes. While the amphibian assemblages are predominantly influenced by the European temperate climatic conditions, the Mediterranean climate plays a key role for the composition of the reptile fauna.

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

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

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

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Species–area relationships and nestedness patterns were studied in three groups of small terrestrial vertebrates (mammals, reptiles, amphibians) on 14 landbridge islands of the eastern Adriatic. islands ranged in surface area between 15 and 410 km2 and contained from eight to 36 species from a total species pool of 48. reptiles were the most species rich group (S = 28), and had more species than mammals (S = 13) and amphibians (S = 7) combined. island surface area predicted species richness best in reptiles (r2 = 0.79) and most poorly in amphibians (r2 = 0.52). Mammals showed a significantly lower slope of the species–area curve than amphibians and reptiles, and thus accumulated species counts with increase in area at the lowest rate. Nestedness patterns in all groups were significantly more organised than expected by chance. Amphibian nested structure points to extinction dominated and well insularised populations with no subsequent recolonisations. frequent unexpected presences and absences in the nestedness patterns of mammals and reptiles suggest complex biogeographic histories for these two groups, with several factors putatively in operation: heterogeneity in habitats and the original source fauna, post- isolation immigrations and differential extinction rate due to human-caused habitat degradation.

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nterspecific aggression is thought to be driven by competition over either shared resources or mates, with the latter facilitated by mistaken or poor species recognition. However, such aggression may potentially also be modulated by other factors, including residency in territorial species. We tested the relative strengths of intra- and interspecific aggression in the lacertid lizard Podarcis melisellensis by introducing males to both the territories of conspecific males and the territories of a sympatric lacertid, Dalmatolacerta oxycephala. We also conducted reciprocal introductions to test the effect of residency on interspecific aggression in P. melisellensis. Our results show that P. melisellensis exhibit significantly more aggression towards D. oxycephala than towards conspecifics, even though these two species do not closely resemble one another and do not exhibit extensive overlap in diet preferences. We also found an overall effect of residency on behavioural measures of aggression, as well as a clear increase in interspecific aggression towards D. oxycephala in resident relative to non-resident P. melisellensis. These results show that interspecific aggression between sympatric species can exist in the absence of breeding competition and with little resource overlap.

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The Sharp-snouted Rock Lizard, Dalmatolacerta oxycephala (Dumeril & Bibron, 1839) is an endemic species of the Eastern Adriatic coast. Distribution area is very narrow, expanding through Southern Croatia, East Hercegovina and Southern Montenegro. As saxicolous species, it is distributed on almost all localities with rocks and cliffs in eumediteranean and submediterranean climate in this region, including most of large and small islands. As very narrow distributed specialist, studies involving this species are very scarce. The goal of this study was to investigate some ecological aspects of this species. Study areas were localized on Island of Vis, Croatia. During four years data on 105 specimens were collected and several ecological parameters of microhabitat were recorded during 24 hours cycles, all year round. Results on refuge selection by night, night thermoregulation and refuges and thermoregulation in winter months were obtained. Also there are some evidences of night activity of this species. Although the Sharp-snouted Rock Lizard was not main target of the research, data collected here can contribute to understand the behavioural patterns of this endemic species.

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The southern part of Montenegro has been identified as an area with high diversity of herpetofauna. However, comprehensive studies of distribution and diversity patterns of reptiles on the country level are still missing. Such studies are essential in designating areas of special conservation importance and nature protection planning in a milieu of increased habitat loss and degradation due to rapid urbanization and tourism development in this small Mediterranean country. To make progress on this problem, we analyzed distribution and diversity patterns of the lizards in Montenegro using a large database consisting of literature data and our unpublished records. We found that fifteen lizard species inhabit Montenegro, and two additional species may be present. The lizards were most diverse in the Maritime biogeographic region of Montenegro, while low diversity was found predominantly along the state borders in the Mountain-valley region. The identified pattern of lizard diversity is at least partly influenced by sampling bias. The eastern mountainous subregion had a distinct species composition compared to all other parts of the country. The East-Mediterranean chorotype was the most dominant, represented by seven species. The great diversity of the lizard fauna of Montenegro can be attributed to its specific topographic position with great influence of Mediterranean climate, heterogeneity of biomes, complex geological history and diverse physiogeographic features. High lizard species richness in the Maritime region and a unique species composition in the eastern subregion of Montenegro indicate that these areas are of high conservation interest.

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Geometric morphometric techniques were used to examine allometric and non-allometric influences on sexual shape dimorphism (SShD) in the ventral cranium (skull base, palate and upper jaw) of four species of lacertid lizards (Podarcis muralis, Podarcis melisellensis, Dalmatolacerta oxycephala, Dinarolacerta mosorensis). These species differ in body shape, ecology and degree of phylogenetic relatedness. The structures of the ventral cranium that were studied are directly involved in the mechanics of feeding and are connected to the jaw musculature; these structures are potentially subject to both sexual and natural selection. Allometry accounted for a considerable degree of cranial shape variation between the sexes. Allometric shape changes between individuals with smaller cranium size and individuals with larger cranium size are mostly related to changes in the skull base showing pronounced negative allometry. The rostral part, however, either scaled isometrically or showed less pronounced negative allometry than the skull base. Non-allometric intersexual shape variation predominantly involved changes related to the jaw adductor muscle chamber, i.e., changes that are associated with biomechanically relevant traits of the jaw system in females and males. Both allometric and non-allometric shape changes appeared to be species-specific. Our results indicate that natural and sexual selection may be involved in the evolution of SShD.

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

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Species composition and abundance of reptiles was examined in three different habitats on the island of Cres (Croatia) adjacent to the town of Osor in May 2013. The aim of our study was to examine the sensitivity of biodiversity and similarity indices in reptiles. In each habitat visual observations of reptiles were made by groups of 10 people for 1 hour along a 1 km sampling path. In the first habitat, overgrown pastures surrounded by stone walls, 5 lizard (Podarcis melisellensis, Algyroides nigropunctatus, Podarcis muralis, Lacerta viridis and Pseudopus apodus) and 2 snake species (Hierophis gemonensis and Elaphe quatuorlineata) were observed. In the second habitat, a gravel road on Osorščica hill surrounded by macchia, 3 lizard (P.melisellensis, A. nigropunctatus and L. viridis) and 2 snake species (H. gemonensis and E. quatuorlineata) were found. In the final habitat, stone buildings and walls in the town, 3 species of lizards (P. melisellensis, A. nigropunctatus and Dalmatolacerta oxycephala) were recorded. The dominant species in all three habitats was the Dalmatian Wall Lizard (P. melisellensis). Two biodiversity indices were applied to the data. Simpson’s diversity index indicated that that the second habitat (0.56) had the greatest diversity in terms of species and their abundance, while the Margalef’s index indicated that the first habitat had more diversity (1.69). In addition, Jaccard’s index of similarity indicated that the urban habitat was dissimilar to the other two, while the Bray-Curtis dissimilarity index showed a weak differentiation between the overgrown pastures and the other habitats. The discrepancy in the outputs from the indices highlights the importance of choosing an appropriate mathematical construct to examine biodiversity and habitat similarities.

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

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A commented lacertids list for Europe, Africa, the Middle East, including the Arabian Peninsula and Asia is given.

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

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

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

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Although Albania has a rich reptile fauna, efforts to reveal its diversity have so far been limited. To fill this gap, we collected available published and unpublished (museum collections, online sources) records of reptile occurrences and conducted several expeditions to search for reptiles in areas with few or no previous records. Our georeferenced database contains 3731 records of 40 species from between 1918 and 2015. Based on this comprehensive dataset, we prepared distribution maps for each reptile species of the country. Applying spatial statistics, we revealed that overall sampling effort was clustered, with hotspots associated with easily accessible areas and natural heritage sites. The maximum number of species per cell was 26 with an average of seven. Cells harbouring large reptile diversity were located along the Adriatic and Ionian coasts, on the western slopes of south Albanian mountains, i.e. in areas generally considered as Balkans biodiversity hotspots or potential historical refugia. We found that species presence and diversity is strongly influenced by landscape features. Diversity of land cover, altitudinal variation, temperature and precipitation variation explained the observed pattern in our models. Our study presents the largest database of reptile occurrences to date and is the first to analyse reptile diversity patterns in Albania. The database and the diversity patterns can provide a basis for future macroecological studies and conservation planning.

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

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

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

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The Balkan Peninsula is a hot spot for European herpetofaunal biodiversity and endemism. The rock climbing lizards Dalmatolacerta oxycephala and Dinarolacerta mosorensis and the ground-dwelling Dalmatian wall lizard Podarcis melisellensis are endemic to the Western Balkans, and their ranges largely overlap. Here, we present a comparative phylogeographical study of these three species in the area of their codistribution in order to determine the level of concordance in their evolutionary patterns. Phylogenetic analyses were performed based on two mitochondrial genes (cytochrome b and 16S rRNA), and a molecular clock approach was used to date the most important events in their evolutionary histories. We also tested for correlations regarding genetic differentiation among populations and their geographical distances. For all three species, a significant correlation between genetic and geographical distances was found. Within D. oxycephala, two deeply separated clades (‘island’ and ‘mainland clade’), with further subdivision of the ‘mainland clade’ into two subclades (‘south-eastern’ and ‘north-western’), were found. High sequence divergences were observed between these groups. From our data, the time of separation of the two main clades of D. oxycephala can be estimated at about 5 mya and at about 0.8 mya for the two subclades of the mainland clade. Within D. mosorensis, coalescence time may be dated at about 1 mya, while D. mosorensis and D. montenegrina separated around 5 mya. The results imply the existence of complex palaeo-biogeographical and geological factors that probably influenced the observed phylogeographical patterns in these lacertid species, and point to the presence of numerous glacial/interglacial refugia. Furthermore, the observed cryptic genetic diversity within the presently monotypic species D. oxycephala prompts for a revision of its taxonomic and conservation status.

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At the western side of Dinarides-Hellenides mountain chain 12 species of lacertid lizards occur. All hitherto investigated species appeared to be phylogeographically highly structured. Their present phylogeographic patterns are the result of the complex topography and the geotectonic history of the region, multiple events of range retractions and subsequent expansions during Pleistocene climatic oscillations but also distinct ecological preferences of the species, interspecific competition, overwater dispersals and unintentional anthropogenic introductions. The phylogeography of Podarcis siculus, P. melisellensis, P. tauricus, Algyroides nigropunctatus, Dalmatolacerta oxycephala and Dinarolacerta mosorensis will be discussed in details.

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This paper presents the data on specimens of genera Algyroides, Dalmatolacerta and Dinarolacerta (Lacertidae) stored in the Herpetological Collection of the Natural History Museum of Montenegro. This part of Herpetological Collection includes 185 specimens belonging to 3 species: Algyroides nigropunctatus (40 specimens), Dalmatolacerta oxycephala (17 specimens) and Dinarolacerta mosorensis (128 specimens). All specimens were collected in Montenegro. The paper provides all relevant data on collected specimens.

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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.

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The present study reports the morphology of leukocytes of 12 European lacertid lizards (Podarcis sicula, P. tiliguerta, P. melisellensis, P. bocagei, P. muralis, Algyroides nigropunctatus, Lacerta viridis, L. bilineata, L. trilineata, L. oxycephala, Timon lepidus, and Zootoca vivipara) stained using May–Grünwald/Giemsa method. The morphology of white blood cells was very similar among species, suggesting a relative morphological uniformity within the lacertid lizards. For six species (i.e. P. sicula, P. tiliguerta, P. melisellensis, P. bocagei, P. muralis, and A. nigropunctatus), we determined the leukocyte differential counts, which may be considered representative of the normal values of the corresponding populations. These results may be useful either in clinical investigation to detect pathologies in wild individuals, as in management and conservation projects to assess the general health conditions of natural wild lizard populations.

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During two short trips to the island Cres in the Kvarner bay in May 2009 and June 2010 the authors observed an introduced population of the Sharp-snouted Rock Lizard (Dalmatolacerta oxycephala) at a wall near the habor of Osor.

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We studied the thermal ecology of the lacertid lizards Lacerta oxycephala and Podarcis melisellensis on the Adriatic island of Vis (Croatia) during summer. These species obviously differ in microhabitat use: L. oxycephala climbs on rocks and stone walls, whereas P. melisellensis is mainly ground-dwelling in vegetation. Since theoretical considerations predict a difference in thermal quality between the species` microhabitats, this system seems to present a good opportunity to test the influence of thermal microhabitat quality on body temperature, thermoregulatory behaviour and evolution of thermal characteristics. Data on thermoregulatory behaviour, body temperatures (T sub(b)) and habitat quality were collected in the field and selected temperatures (T sub(sel)) were estimated in a laboratory thermogradient. Accuracy and effectiveness of thermoregulation were quantified. Thermoregulatory behaviour consisted of timing of activity, selection of places in full sun and near sun-shade transitions, and basking. As predicted, L. oxycephala occupied the microhabitat with the lower thermal quality and had on average a lower T sub(b). However, L. oxycephala also selected lower temperatures in the experimental thermogradient. Thus, if T sub(sel) can be regarded as the thermoregulatory target, both species proved to be accurate and effective thermoregulators. These results corroborate the `labile view` on the evolution of thermal physiology: both L. oxcephala and P. melisellensis appear to be adapted to their respective thermal microhabitat. This is a surprising conclusion, since earlier studies have found the thermal characteristics of Lacertidae to be evolutionarily rigid.

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During nine stays on Mount Biokovo (1762 m) / Central Dalmatia (Croatia), and the adjacent parts of the Cetina valley in the years 1979–1990, 16 reptile species were observed. Together with previous data from literature the list now comprises 21 species. New or more detailed data are given, particularly on the following lizard species: Lacerta trilineata, L. viridis, L. mosorensis, L. oxycephala, Podarcis muralis and P. sicula.

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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.

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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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The sharp snouted rock lizard, Dalmatolacerta oxycephala, is an endemic lizard of the Balkan Peninsula with 70% of its range found in Croatia and Bosnia and Herzegovina (B&H). The herpetological literature surprisingly yields scarce localized data. In this paper we summarize most, if not all, records found in literature, data from museum collections, our own field surveys and records of field researchers with reliable knowledge of D. oxycephala. All literature and new data are plotted and an updated distribution map for Croatia and B&H is given. The sharp snouted rock lizard occurs continuously over the southern areas of the region with new records increasing the known distribution towards the north and into the high mountainous regions. The species occupies a very wide set of habitats, from rocky shrubs at sea level (on islands) up to rocky mountain cliffs and gorges at 1400 m a.s.l. Overall seasonal activity was highest from April to June with strong affiliation to warm and dry habitats. The species has a large range, it is still very abundant and there is no direct evidence of population decline, therefore its IUCN regional status for Croatia should remain least concerned (LC) and near threatened (NT) for B&H.

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Sharp-snouted rock lizard (Lacerta oxycephala, Dumeril & Bibron, 1839) is a stenoendemic species of former Yugoslavia and the Balkan Peninsula (Crnobrnja-Isailović & Džukić, 1997a). It is distributed along the southeastern region of Adriatic coast, including some islands, from Krka River in Dalmacia, to Lake Skadar Region in Montenegro (Radovanović, 1951 Anrold & Burton, 1978; Džukić, 1991; Crnobrnja-Isailović & Džukić, 1997a,b). In FR Yugoslavia, this species is distributed at the Mediterranean and Submediterranean Montenegro (Brelih & Džukić, 1974; Džukić, 1991, 1995 Crnobrnja-Isailović & Džukić, 1997b).

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The present detailed account of the amphibian and reptile distribution in the North Adriatic Island of Cres (Croatia) is chiefly based on literature data supplemented by new observations kindly provided by a number of correspondents or made by the authors during various trips to the island. Dot maps of the species’ distribution on Cres Island are presented. The overview is completed by information on the herpetofauna of 25 neighboring islands in the Kvarner Bay. Lacerta oxycephala DUMÉRIL & BIBRON, 1839 is reported for the first time from Cres Island.

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In 2005, a new lizard species, the sharp-snouted rock lizard (Dalmatolacerta oxycephala) appeared on the island of Cres, probably due to anthropogenic transfer. Before that, only the Dalmatian wall lizard (Podarcis melisellensis) had inhabited the town of Osor, where the new species was discovered. In areas where the two species live sympatrically, P. melisellensis is competitively superior to D. oxycephala; however, in Osor, D. oxycephala appeared in parts of the town where P. melisellensis had been present before. Since 2005, there has been no systematic survey to assess how the distribution of the two species has changed in Osor and, the aim of this study is to investigate how far D. oxycephala has spread and what happened to the distribution of the native P. melisellensis after the new species appeared. Here we show that in 16 years, the native Dalmatian wall lizard was outcompeted by the sharp-snouted rock lizard, and there was no observation of the two species together at the same locations. Despite the new species preferring high stone walls, specimens were also found on shorter stone walls alongside the road towards Lošinj. According to our findings, the further spread of the sharp-snouted rock lizard cannot be excluded in the upcoming years in Cres, which could negatively affect the native Dalmatian wall lizard, but further research is needed to reveal the causes of our observations and to monitor the changes in the distribution of the two species.

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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.

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We tested the hypothesis that an evolutionary trade-off exists between the capacity to run on level terrain and the ability to climb inclined structures in lacertid lizards. Biomechanical and physiological models of lizard locomotor performance suggest that the morphological design requirements of a ground-dwelling vs. scansorial life style are difficult to reconcile. This conflict is thought to preclude simultaneous evolution of maximal locomotor performance on level and inclined terrain. This notion has been corroborated by comparative studies on lizard species from other groups (Anolis, Chamaeleo, Sceloporus), but is not supported by our data on 13 species from the family Lacertidae. We found no indication of a negative association between maximal sprint speed of lizards over a level racetrack (indicative of ground-dwelling locomotor performance), on an inclined stony surface (indicative of climbing performance over rock faces) and inclined mesh surface (indicative of clambering performance among vegetation). Moreover, morphological characteristics associated with fast sprinting capacities (e.g. long hind limbs) apparently enhance, rather than hinder climbing and clambering performance. We conclude that in our sample of lacertid lizards, the evolution of fast sprinting capacity on level terrain has not inflicted major restrictions on climbing and clambering performance.

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The aim of this study was to determine if divergence in habitat use among lacertid lizards is paralleled by morphological differentiation. For 35 lacertid species, we measured body, head and limb dimensions. Habitat use was inferred from the literature: ground-dwelling on open terrain, ground-dwelling in vegetated areas, shrub-climbing, tree-climbing, saxicolous (i.e. rock-climbing). Traditional (i.e. non-phylogenetic) statistical analyses suggest morphological differences among species groups with different habitat use. Ground-dwelling species from open habitats tend to have longer femurs, tibiae and humeri (relative to body length) than other groups. Cursorial (i.e. level-running) species have relatively high heads and trunks compared to climbing species. These differences follow biomechanical predictions and it is tempting to consider tham as adaptations to habitat use. However, phylogenetic analyses of the data fail to establish a clear relationship between habitat use and morphology in the data set considered. There is a weak indication that the differences in head and trunk height have evolved as an adaptation to different habitat use, but the differences in relative limb dimensions among species groups with different habitat use vanish. Either adaptation of limb dimensions to habitat use has not occurred in lacertid lizards, or our methods are unable to demonstrate such an adaptation. We show that uncertainties in the topology of the phylogenetic tree used are unlikely to influence the outcome of our study. We also address the fact that habitat use is often similar in different branches of the phylogenetic tree, and the consequences this may have for the power of our statistical analyses.

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Microhabitat use and antipredator behaviour (i.e. behavioural response to a bird model and refuge microhabitat) was measured in an experimental set-up for 11 lacertid lizard species. Additionally, locomotor performance (sprint speed, climbing and clambering speed, manoeuvrability and endurance) of the same species was quantified. The results showed considerable interspecific variation in microhabitat use, behavioural response and choice of refuge microhabitat. Part of that variation could be linked to differences in locomotor capacity. Species that frequently made use of open microhabitats tended to be fast sprinters, but had limited endurance. The behavioural data suggest these species will not start running before the predator is close at hand, and thus having high sprint capacities might be more beneficial than great endurance. Species that spent most of their time on vertical elements appeared to be fast climbers. As such microhabitats tend to be quite exposed, lizards living there are conspicuous to predators and being able to escape fast seems relevant. Contrary to our expectations, species mostly observed in the vegetated microhabitats did not excel in clambering capacity or manoeuvrability. Possibly, dense vegetation constitutes a safe harbour and species may no longer be under selection for locomotor speed. Behavioural response when confronted with an aerial predator was context-dependent in most species: most lizards fled more when in open or vertical microhabitats than when in densely vegetated ones. Moreover, when fleeing most species sought refuge in the vegetation. These findings support our idea that vegetated patches may constitute a safe harbour for lacertids.

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Morphological and physiological considerations suggest that sprinting ability and endurance capacity put conflicting demands on the design of an animal`s locomotor apparatus and therefore cannot be maximized simultaneously. To test this hypothesis, we correlated size-corrected maximal sprint speed and stamina of 12 species of lacertid lizards. Phylogenetically independent contrasts of sprint speed and stamina showed a significant negative relationship, giving support to the idea of an evolutionary trade-off between the two performance measures. To test the hypothesis that the trade-off is mediated by a conflict in morphological requirements, we correlated both performance traits with snout-vent length, size-corrected estimates of body mass and limb length, and relative hindlimb length (the residuals of the relationship between hind- and forelimb length). Fast-running species had hindlimbs that were long compared to their forelimbs. None of the other size or shape variables showed a significant relationship with speed or endurance. We conclude that the evolution of sprint capacity may be constrained by the need for endurance capacity and vice versa, but the design conflict underlying this trade-off has yet to be identified.

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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.

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This work summarizes the published and unpublished news about the presence in the Italian geographical and political territory of the species of amphibians and reptiles that are not indicated as such in the most recent publications on the subject (Sindaco et al., 2006; Lanza et al., 2007; Lanza et al., 2009; Corti et al., 2010) or which, for editorial reasons, have been summarized (Di Nicola et al., 2019).

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In this paper we examine the distribution of amphibians and reptiles over the islands of the newly founded Lastovo Archipelago Nature Park (Lastovsko oto~je), Croatia. On several field trips between 1996 and 2008, we encountered five species of lizards (Podarcis sicula, P. melisellensis, Dalmatolacerta oxycephala, Hemidactylus turcicus and Pseudopus apodus), one species of snake (Dolichophis caspius), one species of sea turtle (Caretta caretta) and one species of toad (Bufo viridis). We confirm literature data on the presence of some of these species on different islands of the archipelago, and add distributional records for several other islands and islets. Logistic regression analyses show that the variables predicting presence/absence from different islands vary among species. Within species, populations from different islands noticeably vary in body size and shape, scalation, dorsal and ventral coloration, behaviour and density. This observation adds to the value of the study area as a »natural laboratory« for future research into the ecology and evolution of island populations. Although most species seem to be doing well presently, we list a number of possible hazards and concerns.

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

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

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

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1. Relationships between morphology, bite force capacity, prey handling efficiency and trophic niche were explored in two sympatric species of lacertid lizards, Podarcis melisellensis (Braun 1877) and Lacerta oxycephala Duméril & Bibron 1839. 2. Head shape showed little variation, but head size (absolute and relative to snoutvent length, SVL) differed between species and sexes. Males have larger heads than females, both absolute and relative to their SVL. In absolute terms, male P. melisellensis have larger heads than male L. oxycephala, but the reverse case was true for the females. Relative to SVL, L. oxycephala have larger heads than P. melisellensis. 3. Bite force capacity was estimated by having the lizards bite on two metal plates, connected to a piezoelectric force transducer. Differences in maximal bite force between species and sexes paralleled differences in absolute head size. Differences in body size and head size explain the higher bite force of males (compared with females), but not the higher bite force of P. melisellensis (compared with L. oxycephala). Among individual lizards, bite force correlated with body size and head size. 4. Prey handling efficiency, estimated by the time and number of bites needed to subdue a cricket in experimental conditions, also showed intersexual and interspecific variation. This variation corresponded to the differences in maximal bite capacity, suggesting that bite force is a determining factor in prey handling. Among individual lizards, both estimates of handling efficiency correlated with maximal bite force capacity. 5. Faecal pellet analyses suggested that in field conditions, males of both sexes select larger and harder prey than females. There was no difference between the species. The proportion of hard-bodied and large-sized prey items found in a lizard`s faeces correlated positively with its bite force capacity. 6. It is concluded that differences in head and body size, through their effect on bite force capacity, may affect prey selection, either directly, or via handling efficiency.

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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.

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Thirteen reptilian and two amphibian species were observed at Lake Vrana and the adjacent Adriatic coastal area (Dalmacia, Croatia). Lacerta oxycephala was found in Drage, about 30 km north of the known range of the spe- cies.

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CHRISTMAS CHILOREN. PART 2 This arricle is a sequel ro a recenr arricle of rhe aurhor (van WESTBROEK, 200I}, in which rhe reproduceion of some smal! species of European lizards (Podarcis melisellemis, Lncerttt oxycep/;n/n en Alg;lfoides nigropunctntis) is discussed. In rhis parr special arrenrion is given ro rhe incubarion of rhe eggs and rhe rearing of rhe young animals. Differenr subsrrares can be used for rhe incubarion of rhe eggs, bur rhe aurhor prefers porring composr for succulenrs. This has a good airy srrucrure and has good moisr-regularing qualiries. W hen using nor roo high incubarion-remperarures, for insrance 24,5°C, rhe harchlings are somewhar larger, wirh a bigger porrion offemalcs, compared ro when using higher incubarion-remperarures. The juveniles are reared in smal! conrainers, dimensions 45x25x25 cm (lxwxh), wirhour a lid and wirh a smal! spot- light (incandescenr 40 W or halogen 50 W wirhour rhe glass UV-prorecrion). Using plastic terrariums wirhout seams keeps rhe crickers inside. Ar firsr rhe juveniles are fed smal! crickers, unril rhey reach a lengrh ar which rhey can ear buffalo-worms. C rickers and buffi1lo-worms can be fed ground per- food for dogs. In rhe drinking-warer Calcium Iae- tare is dissolved, one reaspoon in a beer-glass. Warer-soluble viramins are added in a concenrra- rion of I mi per lirre water. The vitamin-supple- ment comains 50.000 I.E. vitamin A and I0.000 I.E. vitamin 03 per mi. Ir is viral rhar rhe li7..ards as wel! as rhe insecrs are fed properly and rhar rhe drinking-warer is rene- wed every day. lf you are a bir slack wirh rhis, the resulrs of rearing rhe young lizards wil! decline.

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In this paper we present distributional data for reptiles from different localities in the Western Balkans. During an 11 year period (2002-2012) we collected new data in the frame of spring student research camps organized by the Biology Students’ Society (DŠB). Surveys were conducted by members of the herpetological groups that were mostly biology students. Surveys were located on five islands Dugi otok, Brač, Korčula, Mljet, and Pag, one peninsula Pelješac (Croatia), in the continental area around Kamenovo and Lake Skadar (Montenegro), Dojran Lake (Macedonia), at lower Neretva Delta (Croatia) and around Niš (Serbia). Altogether 34 different reptile species were recorded in the surveys. The species lists comprised of 7-22 different species per single locality. The highest reptile biodiversity was recorded around Dojran Lake in Macedonia.

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In this paper we present an attack of the centipede Scolopendra cingulata Latreille, 1829 on the Sharp-snouted Rock Lizard, Dalmatolacerta oxycephala (Duméril & Bibron,1839). Shortly after the attack, which presumably lead to the lizards’ death, the centipede pulled its prey intoa hiding place under the rock (Fig. 1). This observation was made on July 7, 2014, in Medvinjak on the island of Korčula (Croatia). It was recorded and reported by Ivan Željković. In this case it is assumed that the lizard seriously underestimated the strength of the centipede in any of the two possible options. The possibility of a direct attack on the lizard by the centipede is the most realistic considering the relationship between the sizes of these animals, but the possibility of the lizard’s attack on a centipede and the situation that prey becomes predator also cannot be rejected. Anyway, the lizard underestimated the power of the centipede as in this first case did not result in its escape (illusion of neutralism: lizard considers its relationship with centipede as neutral and does not run away) and the second wrong choice of a prey (paradoxical predation: prey becoming a predator). Third option that the centipede scavenged on the dead corpse was rejected because the lizard was still fresh (weakly movement of the tail and extremities was observed) and because centipedes are predominantly carnivorous (Lewis 2007, Sierwald & Bond 2007). Underestimation of the centipede as the prey was recently reported as failed ingestion of this species by a young sand viper (Arsovski et al.2014). However, in this case both reptile and centipede were found dead and it can be described as illusion of predation. Centipedes of the genus Scolopendra are known as dangerous and aggressive predators and as animals that are extremely difficult to predate on (eg. Dugon & Arthur 2012, Arsovski et al. 2014). It is known that lizards eat centipedes, but notto a great extent (e.g. Hódar 1996). Centipedes need constantly moisture because they lack the waxy cuticle of insects and arachnids (Barnes 1982), so that the encounters of these animals are probably rare. Predation on lizards (and other vertebrates) by arthropods is often noted (Bauer 1990, Schwammer & Baurecht 1988, Blondheim & Werner 1989, Armas 2000, Toledo 2005). Wrong choice of prey, in terms of trying to feed on potentially dangerous prey is also commonly reported (Willson & Hopkins 2011. Jablonski &Vlček 2012. Šukalo et al. 2013).