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

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The east Asian lacertid lizard genus, Takydromus, is well-defined but its relationships to other primitive Palaearctic lacertids are still uncertain and, although it has similarities to the lower Miocene Miolacerta. Takydromus does not appear to be identical with this. The 16 or 17 recognizable species are reviewed and a key provided for their identification; the remaining area of uncertainty about species boundaries is the Takydromus sexlineatus group in Assam. A phylogeny for Takydromus is estimated using a data set of 35 morphological characters drawn from external features, skeleton and soft-parts. There are two main constituent clades which may be informally recognized as Takydromus s.s. and Platyplacopus. Platyplacopus is subtropical and perhaps relict, whereas Takydromus ranges from temperature to tropical areas. The two clades have different ranges of eggs per clutch. In both of them, elongate forms that climb-extensively in flimsy vegetation have evolved and have independently acquired similar features that confer performance advantages in this situation. Interestingly, the order of assembly of these features is markedly different.

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The east Asian lacertid lizard genus,Takydromus, is well-defined but its relationships to other primitive Palaearctic lacertids are still uncertain and, although it has similarities to the lower Miocene Miolacerta, Takydromus does not appear to be identical with this. The 16 or 17 recognizable species are reviewed and a key provided for their identification; the remaining area of uncertainty about species boundaries is the Takydromus sexlineatus group in Assam. A phylogeny forTakydromusis estimated using a data set of 35 morphological characters drawn from external features, skeleton and soft-parts. There are two main constituent clades which may be informally recognized asTakydromus s.s.and Platyplacopus. Platyplacopusis subtropical and perhaps relict, whereasTakydromus ranges from temperate to tropical areas. The two clades have different ranges of eggs per clutch. In both of them, elongate forms that climb extensively in flimsy vegetation have evolved and have independently acquired similar features that confer performance advantages in this situation. Interestingly, the order of assembly of these features is markedly different.

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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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A complete catalogue is provided for the type specimens of lacertid lizards in the herpetological collection of the Zoological Museum, Berlin, as of December, 1993. The collection contains a total of 514 type specimens, representing 63 taxa, of which 42 are currently regarded as valid at the specific or subspecific level. Types representing an additional four taxa appear to have been lost from the collection. The collections are especially rich in African lacertid types and in historically significant specimens collected early in the Nineteenth Century by Pallas, Eversmann, and Hemprich & Ehrenberg. Also present are the type series of many subspecies of Podarcis lilfordi and P. pityusensis described by Eisentraut.

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The spatial structure of a tropical lizard community living in the Cue Phuong National Park (Northern Vietnam) was studied in December 1988, September 1989, and April 1991. The community is made up of 15 species which belong to 5 families: Gekkonidae (3 species), Agamidae (3), Scincidae (7), Lacertidae (1), and Varanidae (1). Habitat partitioning and microhabitat utilization of the lizards were analyzed. The environment at Cue Phuong can be assigned to three major habitat types: tropical forest, grassland, and river banks. 15 kinds of microhabitats were distinguished. Cluster analysis revealed 4 groups of species: inhabitants of big high trees (2 species), `other arboreal` species (5), forest floor dwellers (7), and grassland inhabitants (1). The most abun- dant species at Cue Phuong is Sphenomorphus trilaeniatus, followed by Acanthosaura lepidogaster, which is the most abundant arboreal species. Many species are patchily distributed within the forest.

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Cuc Phuong National Park is located about 100 km southwest of Hanoi within the tropical rain forest zone and occupies 22500 hectares. 24 species of lizards of 6 families known to occur in the Cuc Phuong are listed together with notes on their ecology. The presented material is based on our observations made in Cuc Phuong during some visits from 1988 to 1991. Two species in the list (Scincella reevesii and Tropidophorus hainanus) are recorded the first time in this site.

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Data on lizards of three families (Dibamidae, Lacertidae, and Anguidae) of the Vietnamese fauna, deposited in the herpetological collection of the Zoological Museum of Moscow University, are given. The following data are presented for every sample: accession number, the location of capture, habitat, and collector. The places of capture are shown on a map of the country.

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A new lacertid lizard, Takydromus madaensis sp. nov., from the Ma Da forest, the Dong Nai Province (Southern Vietnam) is described. This species has 4 pairs of chin shields; 3 loreals on each side; 3 pairs of supraoculars on each side; parietals separated by interparietal and occipital ones; an additional shield between the prefrontals; ventrals in six longitudinal rows; there are 17 lamellae beneath the fourth toe. A dichotomous key to four species of the genus Takydromus in Vietnam is provided.

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

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The author describes his observations on the Long-tailed Lizard (Takydromus sexlineatus) on Bali, near the City of Ubud. Some details on its ecology and habitat are highlighted. This diurnal lizard was found in a small patch of grassland. Takydromus sexlineatus was seen basking on dry gras and leaves. It shares his habitat with Eutropis (formerly Mabuya) multifasciata, Gekko gecko and Hemidactylus platyurus. On Bali Takydromus prefers a damp and warm environement.

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From the perspectives of biodiversity conservation and management, there is an urgent need to have at hand current synopses of classification and distributions of species. In this paper, we review and summarize the classifications, Chinese and English names, type specimens, type localities and distributions of China’s lizard fauna to promote scientific exchange and species conservation among relevant people in China and internationally. As of December 31, 2020, a total of 230 species of lizard, belonging to 44 genera and 12 families, have been recorded from China, including 4 invasive species: 2 in the family Gekkonidae, 1 in Iguanidae and one in Dactyloidae. There are 115 endemic species, accounting for 50% of this group. The proportion of endemic species to China was the highest in the family Eublepharidae (84.62%), and the number of endemic species to China was the highest in the family Agamidae (42 species). The species distributions and classification in the “Fauna Sinica (Reptilia 2): Squamata (Lacertilia)” were updated. Among the provincial administrative units, there is a larger number of species in the southern provinces, such as Yunnan, Guangdong, Guangxi and Taiwan. Sichuan, Yunnan, and Taiwan harbour relatively more endemic species than others. A total of 154 species of lizards were first discovered in China, and the type localities of these species are concentrated in Yunnan, Taiwan, Tibet, Xinjiang, and Sichuan.

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Cytochrome b (cyt b) gene complete sequences (1143bp) of Takydromus sylvaticus were sequenced. In order to clarify the phylogenetic position of the Takydromus sylvaticus, we investigated the phylogeny of 15 Takydromus spp. distributed in East-Asia by Maximum Parsimony (MP), Bayesian Inference (BI), and Maximum Likelihood (ML) methods using DNA fragments of cyt b genes. The results supported that the Platyplacopus merged into Takydromus and negated the validity of Platyplacopus. Furthermore, the prediction of tertiary structures of cyt b exhibited the CD loop region contain two short helices forming a hairpin arrangement, namely cd1 and cd2. Thermostability analysis shows that the CD-loop region is unstable thermodynamically and may provide mobility to amino acids located at the heme, and might provide high flexibility to the top of ISP (iron–sulfur protein) and the cavity region of Qo binding site. It suggested that the two short helices of CD loop region of cyt b was a dominating portion for ISP binding site.

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Lizards use chemical cues to locate and identify prey and plant food, assess the nutritional quality of food, and detect plant toxins. Among insectivorous lizards, all actively foraging species studied respond strongly to prey chemicals sampled lingually, but ambush foragers do not. Much recent research has been devoted to assessing differential responses to food and nonfood chemicals (i.e., food-chemical discrimination) by omnivorous and herbivorous species and determining whether correlated evolution has occurred between plant diet and plant-chemical discrimination. We conducted experimental studies of food-chemical discrimination by two species of teiid lizards, the omnivorous Cnemidophorus murinus and the actively foraging insectivorous Ameiva ameiva. The omnivore distinguished both prey and plant chemicals from control substances. The insectivore exhibited prey-chemical, but not plant-chemical, discrimination, as indicated by tongue-flicking and biting. A comparative analysis using concentrated-changes tests showed that correlated evolution has occurred between plant consumption and plant-chemical discrimination in a major lizard taxon, Lacertiformes. These results extend and strengthen previous findings of similar correlated evolution to a new group and add to a growing database indicating that omnivorous lizards use chemical cues to assess both prey and plant foods.

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Sampling environmental chemicals to reveal prey and predators and to provide information about conspecifics is highly developed in lizards. Actively foraging lizards can discriminate between prey chemicals and control stimuli, but ambush foragers do not exhibit prey chemical discrimination. Recent experiments on a few species of herbivorous lizards have also demonstrated an ability to identify plant food chemicals. We studied chemosensory responses to chemicals from prey and palatable plants in two species of actively foraging, insectivorous lizards. Both the lacertid Takydromus sexlineatus and the teiid Cnemidophorus gularis exhibited strong responses to prey chemicals, but not to plant chemicals. These findings increase confidence in the relationship between prey chemical discrimination and foraging mode, which is based on data for very few species per family. They also provide data showing that actively foraging insectivores in two families do not respond strongly to plant cues. Such information is essential for eventual comparative studies of the relationship between plant diet and responses to food chemicals. The traditional method of presenting stimuli by using hand-held cotton swabs worked well for T. sexlineatus but could not be used for C. gularis due to repeated escape attempts. When stimuli were presented to C. gularis on ceramic tiles and no experimenter was visible, the lizards responded readily. Presentation of stimuli on tiles in the absence of a visible experimenter may be a valuable approach to study of food chemical discrimination by active foragers in which antipredatory behavior interferes with responses to swabs.

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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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Nineteen novel records of amphibians and reptiles (eight frogs, seven lizards, and four snakes) from the northwestern portion of the Cardamom Mountains are reported. Six of these, including three undescribed species, are new records for Cambodia. This underscores the importance of the Cardamom Mountains and Cambodia as a whole in biodiversity estimates of Indochina and the extent to which it still remains understudied.

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Residual yolk is assumed to be an important source of energy and nutrients during early life in nonmammalian amniotes. Available data show that the mean size of residual yolk is far smaller in lizards than in turtles, snakes, crocodiles, and birds, raising a question of whether residual yolk is of functional significance in lizards. Here, we compared data from 26 lizard species with those from other nonmammalian amniotes to test the hypothesis that residual yolk is functionally less significant in species producing more fully developed offspring. In our sample, species mean offspring water contents ranged from 73% to 84% of body wet mass; species mean proportions of carcass dry mass, fat-body dry mass, and residual yolk dry mass to offspring dry mass ranged from 84% to 99%, 0% to 5.0%, and 0% to 14.4%, respectively. Lizards are, on average, more fully developed at hatching or birth than snakes, as revealed by the fact that the mean proportion of carcass dry mass to body dry mass and offspring water contents were both higher in lizards than in snakes. We conclude that the functional significance of residual yolk during early life is generally less evident in lizards. Even in the lizards where residual yolk is of potential functional significance, this portion of yolk contributes little, if any, to postembryonic growth. Future work could usefully collect data across a wider spectrum of reptile taxa to establish a precocial–altricial continuum and test the hypothesis that species with a smaller amount of residual yolk are closer to the precocial end of the continuum.

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Coexisting species may experience population and range changes alone or jointly in response to environmental change. Here, we used six climate variables and ten modeling algorithms to predict the distribution of two Takydromus species (T. septentrionalis and T. sexlineatus) in China. We identified the sympatric and allopatric areas by comparing projections between the two species based on habitat suitability under present and future climate scenarios. We constructed the hypervolumes of six climate variables for the two species and then evaluated overlaps between hypervolumes. From this study, we know the following. First, minimum temperature of coldest month contributes the most to the prediction of habitat suitability. Second, habitats suitable for the two species will shift northward in response to climate warming. Third, the range of T. sexlineatus will expand across the four future time intervals before 2,100, namely the 2021–2040, 2041–2060, 2061–2080, and 2081–2100 intervals, under both Shared socioeconomic pathway (SSP) 245 and SSP585 scenarios, and the range of T. septentrionalis will also expand in the future except at the 2081–2100 interval under the SSP585 scenario. Fourth, the sympatric areas will contract or expand under the SSP245 scenario and expand across the four future time intervals before 2,100 under the SSP585 scenario. Fifth, the niche hypervolumes of the two species partially overlapped, and the differences in niche centroid show some degree of niche differentiation between the two species. These results allow to conclude that climate warming will not only drive the northward drift of sympatric areas but also increase the size of these areas if nothing is done to limit the emission of greenhouse gases. Given the existence of hybridization and introgression between T. septentrionalis and T. sexlineatus in the field where they coexist, we also conclude that climate warming will increase chances of hybridization and introgression between the two species.

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The Long-tailed Lizard, Takydromnus sexlineatis (Daudin, 1802)(Reptilia: Lacertidae) is recorded from the Sisaket farming district (near Ubon Ratachathani) during July 2017. Brief notes on its habitat are provided.

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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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Objective: To study the karyotype and G banding pattern of Takydromus sexlineatus meridionalis Gunther. Methods: making specimen slides by general drying in the air, staining the specimen slides with 10% Giemsa solution, and making the G banding pattern by digesting with trypsin. Results: Its karyotpye is 38,ZZ(ZW), and the karyotypeic formula is 2n=16T+22ST; its chromosomes can be divided into 3 groups Every pair of chromosomes have their characteristic G banding pattern. Conclusion: It is a typical species of Scincolacertoidae and a primitive species in Saurta,but it is more evolutional than Takydromus septentrinalis Guenther and Takydromus woiteri.

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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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Morphology is an important factor in locomotion. It may guide the control strategies that an animal or a robot uses for efficient locomotion. Based on our previous work for the modeling of the long-tailed lizard, in this paper we explore the effect of tail loss, a morphological feature that is particularly distinctive in this species. The main aim is to postulate and possibly predict the changes in the locomotor strategies and performance of an amputated long-tailed lizard. For our study we use optimization algorithms and we mainly focus on the results from the standard PSO (particle swarm optimization). Overall the effect of tail loss does not alter much the behavior of the model, both in terms of postural kinematics and speed performance. However some results show particular interest: first the amputated model uses half the power for achieving the same performance as the intact one, second the amputated model uses wider foot placement for the hind limbs and significantly smaller spinal oscillatory amplitudes. These results may predict that an amputated animal will experience stability problems at higher frequencies.

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Morphology is an important factor in locomotion. It may guide the control strategies that an animal or a robot uses for efficient locomotion. In this paper we try to understand the locomotion strategies of a lizard with a distinctive feature, the long-tailed lizard Takydromus sexlineatus. We recorded the performance of real animals in terms of forward speed and then developed a simulation model respecting the morphometric characteristics of long-tailed lizards. We then run systematic tests altering several control parameters of the model. The simulation experiments suggested possible control strategies for effective locomotion given this type of morphology. The experiments were not constrained or guided by any prior knowledge on specific animal angular kinematics. Therefore, the good match between the suggested kinematics for optimal speed and the kinematics of the real animal suggests that our framework is capable of exploring in the future the effects of morphosis on the locomotion strategies of animals, e.g. to perform the same study with shorter or no tail.

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We sequenced the mitochondrial genome of the Western green lizard (Lacerta bilineata) using Illumina technology and additional Sanger sequencing. The assembled 17 086 bp mitogenome had a GC content of 40.32% and consisted of 13 protein-coding genes, 22 tRNA genes, two rRNA genes, and one control region (CR), with a gene order identical to the chordate consensus. In addition, we re-sequenced the mitogenome of the closely related Eastern green lizard L. viridis using the same techniques as for L. bilineata. The mitogenomes of L. bilineata and L. viridis showed a sequence identity of 94.4% and 99.9%, respectively, relative to the previously published L. viridis mitogenome. The phylogenetic reconstruction based on 17 Lacertinae mitogenomes using Anolis carolinensis as the outgroup supported L. bilineata and its sister species L. viridis as distinct lineages.

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Brown and green are the most commonly imitated colors in prey animals because both colors occur in a range of habitats. Many researchers have evaluated survival with respect to background color matching, but the pigment cell mechanisms underlying such coloration are not known. Dorsal coloration of East Asian Takydromus lizards has shifted from green to brown or from brown to green on multiple occasions during the diversification of the genus, thus giving us an opportunity to examine the cellular mechanisms of background color matching. Brown and green skin were found to differ with respect to the morphological characteristics of iridophores, with different thicknesses of the reflecting platelets and the cytoplasmic spacing between platelets, despite a shared vertical arrangement of pigment cells, i.e., xanthophores in the upper layer, iridophores in the middle layer, and melanophores at the bottom of the dermal layer, among the different Takydromus lizards. Iridophores of brown skin reflected longer wavelengths of light than those of green skin, which may be attributed to the thicker platelets and longer distances between platelets in brown skin. We discuss the potential role of genetic and intracellular mechanisms explaining the thickness and orientation of the light-reflecting platelets of iridophores in Takydromus lizards.

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The expansion of oil palm plantations in Indonesia is often blamed for triggering the destruction of natural resources and reducing tropical biodiversity. Herpetofauna is one of biodiversity in oil palm plantations, and information about its species diversity is lim-ited. This study aims to reveal the diversity of Herpetofauna diversity in the oil palm landscape in Central Kalimantan. Field inven-tories were conducted on seven types of land cover: young palm, mature palm oil, old palm oil, High Conservation Value areas, secondary forests, shrubs and smallholder plantations. We used the Visual Encounter Method to find Herpetofauna species in all land cover types. This study revealed that the diversity of Herpetofauna species in PT BLP varies based on the type of land cover. The HCV area and Secondary Forest have the largest number of species (12 species), while secondary Forest has the highest abundance among other types of land cover, followed by old oil palm plantations. The Oil palm plantation landscapes with land cover variations, water availability, and riparian areas can be part of the habitat of several Herpetofauna species and are man-aged for sustainable palm oil management schemes.

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Takydromus Daudin is a group of Lacertidae lizards with slender bodies and long tails. Half of the Takydromus spp. are endemic to islands of eastern Asia aligned along the Pacific margin of the East Asian continent. This feature offers a good opportunity to study the effects of glaciations and land connections on the speciation of East Asian fauna. We reconstructed the molecular phylogeny of Takydromus species via the mitochondrial 12S rRNA gene. Phylogenetic analyses using maximum-parsimony, neighborjoining, and maximum-likelihood options do not support a two-subgenera scheme of Takydromus and Platyplacopus proposed earlier. In contrast, the phylogeny of Takydromus species on islands fits the sequential separation of island groups influenced by changes in sea level. The hypothesis in our prediction supports the process of vicariant speciation and multicolonization of grass lizards on eastern Asian islands. At least two obvious colonization events were followed by vicariance events. Because the molecular clock of the 12 rRNA gene was not rejected in our model test, it is possible to estimate times of speciation events. As the most isolated and basal species compared to other temperate and subtropical species of Takydromus, the separation period of T. smaragdinus in the central Ryukyus is the crucial point in estimating the evolutionary rate. Quaternary-origin or Tertiary-origin models are proposed and discussed

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Many species of lizards use caudal autotomy as a defense strategy to avoid predation, but tail loss entails costs. These topics were studied experimentally in the northern grass lizard, Takydromus septentrionalis. We measured lipids in the three-tail segments removed from each of the 20 experimental lizards (adult females) initially having intact tails to evaluate the effect of tail loss on energy stores; we obtained data on locomotor performance (sprint speed, the maximal length traveled without stopping and the number of stops in the racetrack) for these lizards before and after the tail-removing treatments to evaluate the effect of tail loss on locomotor performance. An independent sample of 20 adult females that retained intact tails was measured for locomotor performance to serve as controls for successive measurements taken for the experimental lizards. The lipids stored in the removed tail was positively correlated with tailbase width when holding the tail length constant, indicating that thicker tails contained more lipids than did thinner tails of the same overall length. Most of the lipids stored in the tail were concentrated in the proximal portion of the tail. Locomotor performance was almost unaffected by tail loss until at least more than 71% of the tail (in length) was lost. Our data show that partial tail loss due to predatory encounters or other factors may not severely affect energy stores and locomotor performance in T. septentrionalis.

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The understanding of life-history responses to increased temperature is helpful for evaluating the potential of species for tackling future climate change. Herein, adult southern grass lizards, Takydromus sexlineatus, were maintained under two thermal regimes simulating current thermal environment and a 4 °C warming scenario to determine the effects of experimental warming on female reproduction and offspring phenotypes. Experimental warming caused females to oviposit earlier and more frequently; however, it did not affect other reproductive traits, including clutch size, egg mass and clutch mass. Accelerated embryonic development and energy accumulation rate might have occurred in warmed females. Maternal warming appeared to increase early embryonic mortality, but did not shift hatchling size and locomotor performance. Embryos of oviparous lizards might be more vulnerable to climate change at early stages than at later stages. The impacts of climate change in oviparous lizards might be adverse in the longer term because of the shift in pre-ovipositional embryo viability, which possibly led to a decreased number of hatchlings.

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Two new grass lizard species, previously confused with Takydromus formosanus, were identified in Taiwan using mitochondrial DNA sequences. In the present study, 40 morphological characters and their taxonomic significances were assessed using principal components analysis and discriminant analysis. The three species can be distinguished by morphology of body sizes, dorsal and caudal scales, and most importantly, by male coloration during the breeding seasons. The present study increases the total number of Takydromus to 19 species, of which 10 are insular endemics, with six only occurring in Taiwan.

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

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

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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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Lacertid lizards use chemical cues emitted by saurophagous snakes to evade predation. Whether these lizards can detect and respond to the chemical cues of predatory mammals has not been studied. As many mammals carry distinct body odours and/or use chemical cues for intraspecific communication, lizards can be expected to use these chemicals as early warning cues. To test this idea, we observed the behaviour of Asian grass lizards (Takydromus sexlineatus) that had been transferred to an unfamiliar test arena containing one of four scent treatments. No particular scent was applied to the arena in the control situation. Diluted aftershave served as a pungency control. In the snake treatment, scent of the Oriental whip snake (Ahaetulla prasina) was applied. We included this treatment to learn how Asian grass lizards react to predator chemical cues. Finally, in the mongoose treatment, the lizards were confronted with scent cues of several small Indian mongooses (Herpestes auropunctatus). Snake scent elicited foot shakes, startles and tail vibrations. These are behaviours that in lacertid lizards are associated with stressful situations such as predatory encounters. Surprisingly, lizards confronted with mongoose scent exhibited none of these stress‐indicating behaviours. In fact, their behaviour did not differ from that of lizards subjected to an odourless control treatment. These results raise concern. Mongooses are rapidly invading ecosystems worldwide. If lizards that have co‐evolved with mongooses are unable to detect these predators’ presence through chemical cues, it seems highly unlikely that evolutionary naïve lizards will develop this ability rapidly.

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The morphology and reproductive biology of Takydromus sexlineatus were studied in Bogor. Males are larger than females. There is sexual dimorphism on head size and tail length (larger and longer tail on males) of body size among sexually mature adults, apparently as a consequence of sexual selection. The diet of T. sexlineatus consists of variety of insects and their larvae, arachnids, decapods and snails. Most of the diet are insect (more than 70 % for adults and juveniles). There are higher prey competitions at adult skinks between males and females, adult females and juveniles, but not between adult males and juveniles. The length of adult males and females are 44.6 - 58.1 mm and 49.6 - 62.3 mm respec- tively, lay eggs throughout the year with 2 -3 clutches, per clutch consists of 1 - 3 eggs.

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As a result of recent field surveys in 2013 and 2018 we herein report the herpetofaunal list of Thai Nguyen Province, comprising 26 species of amphibians (19 genera, seven families, two order) and 72 species of reptiles (51 genera, 20 families, two order). Of which, 17 are threatened species, including eight species listed in the Governmental Decree No. 32/2006/ND-CP, 14 species listed in the Vietnam Red Data Book (2007), seven species listed in the IUCN Red List (2018), and one species listed in the Governmental Decree No. 160/2013/ND-CP. 11 of them, Megophrys major, Occidozyga lima, Odorrana cf. bacboensis, Polypedates mutus, Gehyra mutilata, Lygosoma quadrupes, Varanus salvator, Dendrelaphis pictus, Hebius boulengeri, Xenochrophis trianguligerus, and Sinomicrurus macclellandi are recorded for the first time from this province.

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A new case of female heterogamety has been described in Takydromus sexlineatus. In this species the W-chromosome has the same morphology as the Z-chromosome, but differs from it in being heterochromatic and C-banding positive. This situation is similar to that found in some snakes, and is considered by some investigators as a primitive step in the differentiation of sex chromosomes. This suggests that the mechanisms involved in sex chromosomes differentiation in lizards may be the same as those suggested for snakes.

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The occurrence and form of sex chromosomes were investigated with the aid of C-banding and 4′-6-diamidino-2-phenylindole (DAPI) staining in 13 species of lacertid lizards. The results obtained show the presence in five species of a female heterogamety in which the two sex chromosomes have the same shape and size, but the W differs from the Z in being almost entirely heterochromatic. This condition is clearly similar to that found in some snakes and considered to be an early stage of differentiation of sex chromosomes by Singh et al. (1976, 1980). A more evolved condition may be that found in three other species in which the W is distinctly smaller than the Z. A third situation is that found in all Podarcis species which, even though they are considered to be among the more evolved species in the family, possess two sex chromosomes that are indistinguishable. In general, the situation in lacertids may be compatible with the hypothesis of sex chromosome evolution put forward by Singh et al. (1976, 1980). However a differentiation mechanism of this kind does not seem to be well established in lacertids, and is probably not the only mechanism that is in operation in this family.

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Chromosomal inter- and intraspecific variation in Lacertidae. The karyology of various species from the family Lacertidae (Reptilia, Sauria) has been studied with conventional and C-banding techniques. The study shows that this family is not so conservative from a karyological viewpoint as considered til! now. In fact supernumerary chromosomes have been observed in two of the species investigated. Moreover an inter- and intraspecific variability has been found in the C-banding pattern of the various species studied. The situation found in Podarcis sicula is particularly remarkable. Different Cbanding patterns have been observed in three different subspecies and in two populations of the same subspecies. The C-banding technique evidences also the existence in some of the species studied of a female sex heteromorphism, in which the W chromosomes is completely heterochromatic. These observations suggest that in lacertid lizards the evolution of the sex chromosomes follows the same steps hypothesized for snakes.

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The karyology of various species from the family Lacertidae (Reptilia, Sauria) has been studied with conventional and C-banding techniques. The study shows that this family is not so conservative from a karyological viewpoint as considered till now. In fact a higher diploid number than that generally observed in most species of Lacertidae and supernumerary chromosomes have been observed in three of the species investigated. Moreover an evident interand intraspecific variability has been found in the C-banding pattern of the various species studied. The situation found in Podarcis sicula is particularly remarkable. Different C-banding patterns have been observed in three different subspecies and in two populations of the same subspecies. These variations do not show a well defined trend and their role in the evolution of lacertid lizards is discussed. The C-banding analysis evidences also the existence in two of the species studied of a female sex heteromorphism, in which the W chromosome has the same shape and size as the Z, but differs from it in being completely heterochromatic. This situation reminds that observed in some snakes and suggests that in lacertid lizards the evolution of sex chromosomes might have followed the same steps previously hypothesized for ophidians.

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Research of lizards (Sauria) in the Harapan Rainforest, Jambi was conducted from February to April 2012. The research was conducted using the direct capture and survey method. The result found of 14 species of lizards belongs to five families, which are Draco sumatrana, Aphaniotis fusca, and Bronchocela cristatella (Agamidae), Varanus salvator and Varanus sp. (Varanidae), Eutropis multifasciata and Eutropis macularia (Scincidae), Takydromus sexlineatus (Lacertidae), Cyrtodactylus sp., Hemidactylus platyurus, Hemidactylus frenatus, Hemidactylus sp., Gekko smithii and Gekko monarchus (Gekkonidae).

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Historical relationships were inferred for the oriental lizards of the genus Takydromus Daudin 1802 (Lacertidae) on the basis of DNA sequences. Of the 17 species currently recognized for the genus, 13 species represented by 42 specimens from 29 localities were examined. Maximum-likelihood and maximum-parsimony analyses of data for 829 aligned sites from parts of the mitochondrial 12S and 16S rRNA genes yielded relationships that, while showing no substantial discrepancy with each other, were strikingly different from a currently prevailing phylogenetic hypothesis from a parsimony analysis of morphological characters. Based on the results of these molecular analyses, supplemented by results of the morphological analysis that offered robust evidence for positions of two additional species (T. khasiensis and T. sylvaticus), the following interrelationships were hypothesized as the most preferred phylogeny: (kuehnei (sexlineatus khasiensis))(tachydromoides ((smaragdinus (sauteri (dorsalis sylvaticus))) (amurensis (((formosanus wolteri) hsuehshanensis)(toyamai (septentrionalis stejnegeri)))))). These interrelationships indicate: (1) invalidity of Platyplacopus Boulenger 1917, which was recently resurrected as a subgenus of Takydromus on the basis of the morphological analysis; (2) homoplasy in states of some morphological characters, such as green dorsal coloration, that were considered as synapomorphs of certain nodes in the morphological analysis; and (3) involvement of the genus in a series of vicariances in both the continental and insular parts of eastern Eurasia. Due to the paucity of available samples, phylogenetic status of T. intermedius and T. haughtonianus remain to be examined in future studies.

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This study determined the standardized karyotype and idiogram of Takydromus sexlineatus from Surin Province in northeastern Thailand. Mitotic metaphase chromosomes from bone marrow cells and different meiotic stage from testicular cells respectively were studied after staining with conventional Giemsa and Ag-NOR banding techniques for chromosome identification. The study was carried out with samples taken from both male and female T. sexlineatus, the sex of which was identified by external examination. The results showed that the diploid chromosome number of T. sexlineatus was 2n = 38 and the fundamental number (NF) was 38 in both male and female. Although chromosome study was done in both sexes of T. sexlineatus, this study does not identify the sex chromosomes. This is because of the limitation of the staining techniques used. Therefore, we did not conclude on the sex chromosome system in this species and all chromosomes were taken as autosomes. The types of autosomes observed were 16 large telocentric, 14 medium telocentric, and 6 small telocentric macrochromosomes and 2 microchromosomes. Our result is the new record in Thailand on this species and NOR characteristics by silver staining is the first report on NOR banding in this species. The karyotype formula of T. sexlineatus thus was found as: 2n(38) = L 16 t  + M 14 t  + S 6 t  + 2 microchromosomes.

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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 complete sequence mitochondrial genome of Takydromus sexlineatus was determined using long PCR and conserved primers walking approaches. The genome was 18,943 bp in length and contained 13 protein-coding genes, 2 rRNA genes, 22 tRNA genes and 1 control region (CR). The gene composition and order of T. sexlineatus were similar to most other squamate reptiles. All protein-coding genes begin with ATG as initiation codon except COI using GTG. Seven genes (ATP8. ND4L. ND5. Cytb. ND1. COI and ND6) ended with TAA, TAG, AGGA and AGA stop codon, the remaining 6 genes had incomplete stop codons T/TA. The overall base composition of the genome in descending order was 31.48% A, 24.67% C, 30.79% T and 13.05% G, with a slight A + T bias of 62.27%. CR is located between the tRNA-Pro and tRNA-Phe genes and is 3562 bp in length, some tandem repeat sequences, conserved elements (CSB1-3) and termination associated sequences (TAS1-3) were found in the control region.

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The gonads are specific organs since the testes and ovaries, as organs with totally different structure, develop from common, undifferentiated anlages termed genital ridges. Cells sharing the same origin commit various fate, and form different structures, depending on sex determination. The testes and ovaries are therefore a valuable object of research on cell differentiation and mechanisms of organ structure patterning. The development of gonads in reptiles has been studied in few species and many aspects of gonadogenesis in this group of vertebrates have remained unclear. Comparative descriptions showing similarities and differences in gonad development between lizard species have been scarce. The aim of the study was to investigate changes in the structure of developing gonads occurring at the early stages of development in selected species of lizards. The study was carried out on six species of lizards: Correlophus ciliatus, Lepidodactylus lugubris, Eublepharis macularius, Takydromus sexlineatus, Anolis carolinensis and Chamaeleo calyptratus. Gonads were subjected to trichromatic staining specifically revealing connective tissue and basement membranes, which enabled to image the internal structure of the organ. In differentiating testes, the cortex reduces to a thin epithelium, and the germ cells disappear in the reducing cortex, while in the medulla the epithalial cells differentiate and testis cords, containing the germ cells, form. In differentiating ovaries, the cortex grows, which is accompanied by an increase in the number of germ cells in this part of the gonad; the ovarian cortex surrounds the entire medulla or forms only a cluster of cells at the distal gonad pole as in geckos; in the ovarian medulla, cells can eventually differentiate in epithelial structures, and in the leopard gecko even form well-organized cords. The main interspecific differences in the structure of developing gonads concerned a shape of the gonads, their cortex and medulla, and the tendency of ovarian medulla cells to form epithelial structures and cords. The veiled chameleon turned out to be a unique species, in which gonadal development rate is accelerated in relation to soma development. Despite describing significant diversity in the structure of developing lizard gonads, this study reveals the existence of a common model of sexual differentiation of gonads independent of the genetic or temperature sex determination.

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Sex chromosomes are believed to be stable in endotherms, but young and evolutionary unstable in most ectothermic vertebrates. Within lacertids, the widely radiated lizard group, sex chromosomes have been reported to vary in morphology and heterochroma- tinization, which may suggest turnovers during the evolution of the group. We compared the partial gene content of the Z-specific part of sex chromosomes across major lineages of lacertids and discovered a strong evolutionary stability of sex chromosomes. We can conclude that the common ancestor of lacertids, living around 70million years ago (Mya), already had the same highly differentiated sex chromosomes. Molecular data demonstrating an evolutionary conservation of sex chro- mosomes have also been documented for iguanas and caenophidian snakes. It seems that differences in the evolutionary conservation of sex chromosomes in vertebrates do not reflect the distinction between endotherms and ectotherms, but rather between amniotes and anamniotes, or generally, the differences in the life history of particular lineages.

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Among amniotes, squamate reptiles are especially variable in their mechanisms of sex determination; however, based largely on cytogenetic data, some lineages possess highly evolutionary stable sex chromosomes. The still very limited knowledge of the genetic content of squamate sex chromosomes precludes a reliable reconstruction of the evolutionary history of sex determination in this group and consequently in all amniotes. Female heterogamety with a degenerated W chromosome typifies the lizards of the family Lacertidae, the widely distributed Old World clade including several hundreds of species. From the liver transcriptome of the lacertid Takydromus sexlineatus female, we selected candidates for Z-specific genes as the loci lacking single-nucleotide polymorphisms. We validated the candidate genes through the comparison of the copy numbers in the female and male genomes of T. sexlineatus and another lacertid species, Lacerta agilis, by quantitative PCR that also proved to be a reliable technique for the molecular sexing of the studied species. We suggest that this novel approach is effective for the detection of Z-specific and X-specific genes in lineages with degenerated W, respectively Y chromosomes. The analyzed gene content of the Z chromosome revealed that lacertid sex chromosomes are not homologous with those of other reptiles including birds, but instead the genes have orthologs in the X-conserved region shared by viviparous mammals. It is possible that this part of the vertebrate genome was independently co-opted for the function of sex chromosomes in viviparous mammals and lacertids because of its content of genes involved in gonad differentiation.

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We describe a collection of 78 species of amphibians and reptiles from the Cardamom Mountains, southwestern Cambodia. One frog (Calluella guttulata), six lizards (Draco taeniopterus, Dasia olivacea, Lygosoma bowringii, Scincella melanosticta, Sphenomorphus stellatum, and Ptychozoon lionotum), and four snakes (Boiga dendrophila, B. multomaculata, Rhabdophis nigrocinctus, and Xenochrophis trianguligerus) are reported from Cambodia for the first time. Anthopogenically modified environments contain mostly species having broad geographic ranges in Southeast Asia. However, the frog and lizard faunas of intact environments in the Cardamom Mountains are largely distinct from those in the mountainous areas of eastern Cambodia.

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The first amphibian and reptile collection is reported from hilly eastern Cambodia since Henri Mouhot’s work in 1859. The collection contains 30 species of amphibians and 42 species of reptiles. Leptobrachium mouhoti, new species, and Ophryophryne synoria, new species, are described. Of the remaining collection, 11 species of amphibians (39.3%) and seven species of snakes (33.3%) are reported from Cambodia for the first time. Comparisons of the frogs show strong faunal overlap with that of mountainous central Vietnam, but little faunal overlap with the Cardamom Mountains of southwestern Cambodia.

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Patterns in functional diversity of organisms at large spatial scales can provide insight into possible responses to future climate change, but it remains a challenge to link large-scale patterns at the population or species level to their underlying physiological mechanisms at the individual level. The climate variability hypothesis predicts that temperate ectotherms will be less vulnerable to climate warming than tropical ectotherms, due to their superior acclimatization capacity. However, metabolic acclimatization occurs over multiple levels, from the enzyme and cellular level, through organ systems, to whole-organism metabolic rate (hereafter biological hierarchy). Previous studies have focused on one or a few levels of the biological hierarchy, leaving us without a general understanding of how metabolic acclimatization might differ between tropical and temperate species. Here, we investigate thermal acclimation of three species of Takydromus lizards distributed along a broad latitudinal gradient in China, by studying metabolic modifications at the level of the whole organism, organ, mitochondria, metabolome, and proteome. As predicted by the climate variability hypothesis, the two temperate species T. septentrionalis and T. wolteri had an enhanced acclimation response at the whole organism level compared to the tropical species T. sexlineatus, as measured by respiratory gas exchange rates. However, the mechanisms by which whole organism performance was modified was strikingly different in the two temperate species: widespread T. septentrionalis modified organ sizes, while the narrowly distributed T. wolteri relied on mitochondrial, proteomic and metabolomic regulation. We suggest that these two mechanisms of thermal acclimatization may represent general strategies used by ectotherms, with distinct ecological costs and benefits. Lacking either of these mechanisms of thermal acclimatization capacity, the tropical species is likely to have increased vulnerability to climate change.

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Eimeria takydromi n. sp., a coccidian infecting the gall-bladder epithelium and bile-ducts, was found to parasitise three Takydromus species: T. tachydromoides of Honshu, Japan; T. smaragdinus of Amami Oshima, Ryukyu Islands, Japan; and T. sexlineatus of Thailand. Although differences were found in the mean oöcyst and sporocyst indices among the different hosts, the considerable overlap in morphometric characters demonstrated conspecificity. Eimeria takydromi is an addition to the symbiotic complex associated with Takydromus spp. in eastern and southeastern Asia that includes Plasmodium sasai, Trypanosoma takydromi, Schellackia sp. and Lizard Erythrocytic Virus. These host-symbiote associations may have persisted since a lacertid ancestral to the modern species of Takydromus dispersed throughout the region from the late Pliocene.

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A population of Takydromus sexlineatus sexlineatus was observed near Bogor (West Java). A total of 70 lizards were caught after they went to sleep shortly before sunset. Measurements taken revealed no sex-specific proportions but distinctly smaller TL:SVL ratios in juveniles as compared to adults. Nearly all males were found to have ocelli.

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Locomotion is important to animals because it has direct implications for fitness through its role in predator escape, prey capture, and territory defence. Despite significant advances in our understanding of animal locomotion, studies exploring how substrate properties affect locomotor performance remain scant. In the present study, we explore how variation in substrate (sand, slate, cork) affects locomotor performance in lacertid lizards that differ in morphology. Moreover, we explore whether substrate effects are the same for different types of locomotor performance (speed, acceleration, and stamina). Our results show that the substrate affected most types of locomotor performance studied but not always in the same way. Although substrate effects were species-dependent for the maximal speed over 50 cm and the distance run to exhaustion, this was not the case for acceleration capacity. These results suggest that substrate texture differentially affects burst performance vs. longer duration measures of locomotor performance. Finally, straightforward relationships between habitat use and the substrate on which performance was maximized were not observed. This suggests that the evolution of locomotor capacity is complex and that animals may show compromise phenotypes allowing them to deal with a variety of substrates in their natural environment.

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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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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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Over the past two decades many checklists of reptiles of India and adjacent countries have been published. These publications have furthered the growth of knowledge on systematics, distribution and biogeography of Indian reptiles, and the field of herpetology in India in general. However, the reporting format of most such checklists of Indian reptiles does not provide a basis for direct verification of the information presented. As a result, mistakes in the inclusion and omission of species have been perpetuated and the exact number of reptile species reported from India still remains unclear. A verification of the current listings based on distributional records and review of published checklists revealed that 199 species of lizards (Reptilia: Sauria) are currently validly reported on the basis of distributional records within the boundaries of India. Seventeen other lizard species have erroneously been included in earlier checklists of Indian reptiles. Omissions of species by these checklists have been even more numerous than erroneous inclusions. In this paper, I present a plea to report species lists as annotated checklists which corroborate the inclusion and omission of species by providing valid source references or notes.

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

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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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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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This paper deals with habits and characteristics of ecology of Takydromus sexlineatus in the Beiling Mountain in Zhaoqing.The results are as follows(in aresage):The adult body length is 245.104 mm.The tail is 3.9 times as long as the body.The regenerate speed of severed tail is strong.Hibetnation is from December to next February,and below 14℃ they fall asleep completely,while above 20℃ they awake in succession.The most active period is from March to May.Sexual mature females lay multiple clutches of pliable-shelled eggs with clutch sizes of 2~4,and weight of 0.1773 g/egg,long diameter of 8.855 mm and short diameter of 6.157 mm.The incubation needs about 40 days,and the eggs will be hatched in May.

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This report deals with diversity research within the Vietnam project of the Cologne Zoo on his way towards a centre for nature conservation. Since more than five years the Cologne Zoo engages in a German-Vietnamese cooperation project concerning biodiversity research and conservation in the Phong Nha-Ke Bang National Park in central Vietnam. This karst forest area in the Quang Binh province, which meanwhile is declared by the UNESCO as world heritage site, comprises more than 85,000 ha and represents habitat and refuge of many threatened and in part still undiscovered species. My report comprises the current knowledge about the amphibians and reptiles of this area. The so-called herpetofauna appears, due to its limited tendency to spread, the relatively high habitat requirements, and the relatively simple recording (without using traps), as suitable bioindicator. In addition, herpetodiversity research is not only required in Vietnam because being a hot spot (i. e., endemism centre) of biodiversity, but also due to the fact, that we are only able to protect those species which are known so far. In this context, the special significance of newly discovered species as role models for a nature reserve (i. e., so-called flagship species) is stressed: As ambassadors of their relatives they do not only plead for the conservation status of an area, but also motivate the local people to engage in the conservation of their unique home; finally, newly discovered species help to be effective as publicity and to attract support by sponsors, too. Beside newly discovered mammal species some recent reptile discoveries from the National Park’s karst forest are highlighted: Among them is the Phong Nha-Ke Bang Bent-toed Gecko (Cyrtodactylus phongnhakebangensis) that was named after its type locality, as well as another, greyish gecko species (Gekko sp. n.), that will be described as new species soon. However, the area’s most spectacular herpetological discoveries were two horned pitvipers: One of them (Triceratolepidophis sieversorum) was originally discovered as preserved specimen in the rice wine collection of a local healer and revealed not only to be a new species but also to represent a new snake genus; the other horned pitviper (Protobothrops cornutus) has already been regarded by some scientists as being extinct and therefore could be rediscovered in Vietnam after more than half a century (which was at the same time the first record for the National Park’s herpetofauna). Thus, both geckonid as well as both horned pitviper discoveries represent prominent advocates of the habitat karst forest in the Phong Nha-Ke Bang National Park, with whom their existence is closely connected. After these examples, the remaining members of the herpetofauna are briefly introduced in words and pictures: Currently seven families of frogs (represented by almost 40 species), four families of turtles (more than ten species), five lizard families (more than 30 species) and six families of snakes (almost 50 species) are known. Because several problems concerning determination and taxonomy remain still unsolved and because new species are constantly discovered and described, it is difficult to name concrete species numbers. However, with respect to its herpetodiversity, the Phong Nha- Ke Bang National Park meanwhile surely belongs to Vietnam’s best researched nature reserves. Almost 20% of its amphibian and reptile species are listed in the Red Data List of Vietnam. In future, we will engage in proceeding our diversity studies in close cooperation with our Vietnamese partners to provide the National Park as well as the local authorities the knowledge and the required tools for a long-term conservation of the region’s unique biodiversity.