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An annotated list of amphibian and reptile species of the Russian Far East is presented with due account of modern data on the taxa systematics. 13 Amphibian species and 27 reptile ones have been reliably registered by now, including several random sea species.

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Morphologic diversity was studied in 129 species of lacertid lizards and their relationship with ecology by means of comparative analysis on seven linear morphometric measurements. Body size is the most important variable determining, a continuum among small bodied species and larger ones independently evolved through the lacertid phylogeny. This variable is strongly and positively correlated with the others masking the patterns of morphologic diversity. Multivariate analysis on size-adjusted variables show a negative covariation among relative tail and limb length. Remarkably, arboreal and semiarboreal species (Takydromus and the Equatorial African clade) appeared two times independently during the evolution of lacertids and are characterized by extremely long tails, and relatively long forelimbs in comparison with hindlimb length. The arboreal and glider lizard Holaspis with their short tail constitute the only exception. Another case of convergence is found by some species dwelling into dense vegetation or grass (Tropidosaura, Lacerta agilis, Takydromus amurensis or Zootoca) which have long tails and short limbs. On the opposite, species living in deserts, steppes or scrublands with scarce vegetation isolated into extended open areas have developed long hindlimbs and short forelimbs to achieve higher speed and maneuverability This is especially the case of Acanthodactylus and Eremias.

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TERRA TYPICA. Amur region. DISTRIBUTION. It occurs in north-eastern China, Korea and Japan (the island Zussima). In North Eurasia it lives in Russian Far East – in the southern regions of the Primorsky and Khabarovsk Territories. CONSERVATION STATUS. The species does not require special measures on its protection.

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TERRA TYPICA. Chemul’po, Korea. DISTRIBUTION. It occurs in Korea, south-east Manchuria and eastern China from where it penetrates into the south of the Primorsky Territory approximately to the valley of the river Iman in Russia. CONSERVATION STATUS. Not determined.

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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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Relationships of lacertid lizards were assessed on the basis of 84 primary and 112 binary characters drawn mainly from morphology, including features of the skeleton, external anatomy, various internal soft part systems and two aspects of behaviour. Among features not previously used, or not fully investigated before, are structure of the septomaxilla and nasal passages, arranged of the xiphisternal cartilages, mite pockets, kidney position, ulnar nerve arragement, thoracic fascia, aspects of the hemipenis and its associated muscles, female genitalia and jaw muscles. On the basis of parsimony analysis and compatibilty treatment of this character set, the Lacertidae fall into two main portions: A paraphyletic Palaearctic and Oriental group of primitive forms, from which is derived a holophyletic assemblage of Ethiopian and advanced Saharan and Eurasian taxa. The former group ist not fully resolvable, but Psammodromus and Gallotia appear to be sister groups and are probably related to Lacerta parva and L. fraasi and then L. brandtii, Podarcis appears to be related successively to L. andreanszkyi, the sister species L. dugesii and L. perspicillata, and perhaps L. danfordi and L. laevis. This assemblage may be related to archaeolacertas and Algyroides. The separation of Lacerta lepida, L. pater and L. princeps from the agilis group, based on chemical evidence, is weakly contradicted by morphology. Takydromus may be most closely related to L. vivipara, and L. jayakari and L. cyanura constitute the most likely sister group of the Ethiopian and advanced Saharo-Eurasian assemblage. Taxe in the Ethiopian and advanced SaharoEuroasian assemblage form a long essentially pectinate tree with relatively change between the side branches, except for a strong disjunction separating the more primitive from the more advanced taxa. Most of the former fall on two main branches, with ´Lacerta` australis and ´L.` rupicola possibly basal to them. 1. the Equatorial forest group containing Gastropholis, Bedriagaia, ´Lacerta` echinata, Adolfus, ´Lacerta` jacksoni and Holaspis. The first three of these constitute a holophyletic group and the same is probably true of the remainder. 2. Tropidosaura, Poromera and Nucras, the latter being the sister group of the more advanced forms. These include successively the Ethiopian Philochortus, Latastia, Ichnotropis and Heliobolus, Pseuderemias, Meroles and Aporosaura, and Pedioplanis, and then the Saharo-Eurasian Eremias, Acanthodactylus, Mesalina and Ophisops-Cabrita. It seems probable that the ancestors of modern Lacertidae arose in western Eurasia, where the family is known since the Palaeocene and is still represented there largely by quite primitive forms (89 species and seven nominal genera). The family later invaded Africa, perhaps first in the early or middle Miocene. Relatively primitive lacertids spread widely in largely mesic situations in the Ethiopian region, radiating to some extent (six present genera and 16 species) and producing Nucras and the related series of advaned groups (eight genera and 54 species) whoich show increasing adaptation to xeric environments. These genera tend to have heir most primitive species in the northeast and north of the Ethiopian region. The most advaned gave rise to the Saharo-Eurasian clade, now made up to Eremias, Acanthodactylus, Mesalina and Ophisops-Cabrita. This invaded the arid areas of North Africa and Eurasia, where it is presently represented by 70 species. Many morphological changes in increasingly advanced lacertids may be functionally related to the problems of survival in arid, hot, open environments. Considerable ecological parallelism exists in lacertids, with members of separate stocks occupying similar niches in different geographical areas. Morphological adaptations associated with these niches contribute significantly to the high levels of character homoplasy found in the family. There is also some correlation between the degree of niche differentiation in various groups and the quality of the phylogenies that can be produced from their physical characters. A number of morphological parallels exist between advaned lacertids and New World macroteiids. In the skull at least, advaned lacertids show a complex mixture of paedomorphosis and acceleration. Nomenclatorial changes are as follows: Cabrita is synonymised with Ophisops, necessitating a new name, Ophisops nictans, for Cabrita jerdonii. Aporosaura is synonymised with Meroles, Platyplacopus with Takydromus, and Bedriagaia with Gastropholis. ´Lacerta` (or Centromastyx) echinata is also transferred to the latter genus and Lacerta jacksoni to Adolfus. ´Lacerta` australis and ´L.` rupicola are put in a new genus, Australolacerta. It is recommended that Lacerta dugesii and L. perspicillata should not be placed in the otherwise very uniform genus Podarcis. Although clearly paraphyletic, Lacerta s. lat. Should be retained at least for the present and, if necessary putative relationships within it indicated by informal groups or subgenera.

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

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The sclerotic ring consists of several bones that form in the sclera of many reptiles. This element has not been well studied in squamates, a diverse order of reptiles with a rich fossil record but debated phylogeny. Squamates inhabit many environments, display a range of behaviours, and have evolved several different body plans. Most importantly, many species have secondarily lost their sclerotic rings. This research investigates the presence of sclerotic rings in squamates and traces the lineage of these bones across evolutionary time. We compiled a database on the presence/absence of the sclerotic ring in extinct and extant squamates and investigated the evolutionary history of the sclerotic ring and how its presence/absence and morphology is correlated with environment and behaviour within this clade. Of the 400 extant species examined (59 families, 214 genera), 69% have a sclerotic ring. Those species that do not are within Serpentes, Amphisbaenia, and Dibamidae. We find that three independent losses of the sclerotic ring in squamates are supported when considering both evolutionary and developmental evidence. We also show that squamate species that lack, or have a reduced, sclerotic ring, are fossorial and headfirst burrowers. Our dataset is the largest squamate dataset with measurements of sclerotic rings, and supports previous findings that size of the ring is related to both environment occupied and behaviour. Specifically, scotopic species tend to have both larger inner and outer sclerotic ring apertures, resulting in a narrower ring of bone than those found in photopic species. Non-fossorial species also have a larger sclerotic ring than fossorial species. This research expands our knowledge of these fascinating bones; with further phylogenetic analyses scleral ossicles could become an extremely useful character trait for inferring the behaviour of fossil squamates.

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During a trip to Taiwan in May 2018, an Austrian team of herpoetologists recorded three different species of lizards in various habitats. While the mountain dweller Takydromus hsuehshanensis was found in gravel slopes and dump sites above 1.800 m, T. kuehnei was recorded in small clearings and forest margins in the transition zone between planes and hills. Finally T. viridipunctatus was found in gardens and parks in the North of the country near the coast.

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

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Eremias guttulata, Eremias velox, Eremias strauchi, Eremias persica, Eremias regeli, Eremias multiocellata, Eremias nikolskii, Eremias pleskei, Eremias lineolata, Eremias scripta, Eremias intermedia, Eremias nigrocellata, Eremias arguta, Eremias argus, Eremias brenchleyi, Eremias przewalskii, Eremias grammica, Lacerta strigata, Lacerta viridis, Lacerta trilineata, Lacerta agilis, Lacerta parva, Lacerta vivipara, Lacerta taurica, Lacerta brandtii, Lacerta derjugini, Lacerta praticola, Lacerta chlorogaster, Lacerta saxicola, Lacerta caucasica, Lacerta rudis, Ophisops elegans, Takydromus amurensis, Takydromus wolteri.

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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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Takydromus sikkimensis was named in 1888 by A. Günther on the basis of an 1872 description of grass lizards from Sikkim by F. Stoliczka. Subsequent workers have not considered this species valid and have even doubted the occurrence of Takydromus in Sikkim. We confirm the occurrence of grass lizards in the lower Teesta valley of Sikkim. These are consistent with the description of T. sikkimensis and are distinguishable from all other congeners on the basis of having 12 rows of enlarged ventral scales across midbody, three pairs of chin shields, 3–6 femoral pores on each thigh, and color pattern lacking stripes, spots, or ocelli. Because the type specimens are lost, and because there has been a persistent confusion regarding the identity, and even the existence, of the taxon described by A. Günther as T. sikkimensis, we designate a neotype to stabilize the use of the revalidated name.

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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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In the herpetological collection of ZFMK 528 scientific species group names are represented by type materi- al. Of these, 304 names are documented by primary type specimens (onomatophores) while for 224 further names sec- ondary type specimens (typoids) are available, ranging chronologically from 1801 to 2010. The list is a shortened pred- ecessor of a comprehensive type catalogue in progress. It lists name bearing types with their catalogue numbers includ- ing information on further type series members also in other institutions, while secondary types are listed only by pres- ence, both in ZFMK and other collections including holotype repositories. Geographic origin and currently valid names are also 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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First experience with Takydromus septentrionalis (GÜNTHER, 1864) A description is given of Takydromus septentrionalis. Because these animals are potentially suited to keep in an outdoor-terrarium, the climate of their area of distriburion is compared to that of the Netherlands. The husbandry of the animals, partly in an outdoor-terrarium, is discussed. The animals are kept inside during the winter.

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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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Helminths from Eumeces latiscutatus (n = 17) and Takydromus tachydromoides (n = 34) from Japan are reported. Two species of Nematoda, Kurilonema markovi and Meteterakis japonica, and 1 taxon of Acanthocephala, cystacanths of Centrorhynchus sp., were found in E. latiscutatus; 1 species of Digenea, Plagiorchis taiwanensis, and 2 species of Nematoda, K. markovi and Oswaldocruzia filiformis, were found in T. tachydromoides. Oswaldocruzia yezoensis is proposed as a synonym of Oswaldocruzia filiformis. Takydromus tachydromoides represents a new host record for P. taiwanensis and Japan is a new locality record.

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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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We sequenced partial mitochondrial DNA from the cytochrome b gene (1143 bps) for 385 northern grass lizards (Takydromus septentrionalis) from 14 mainland and 14 island populations covering almost the lizard’s entire range to examine the influence of geographic barriers (mountain ranges and water bodies) on the diversification of lineages. Phylogenetic analyses revealed a detailed distribution of three evolutionary lineages (W, E and G). Lineage G included individuals exclusively from Guiyang, in the south-western distributional limit. Lineage W included individuals from the central and western parts of the lizard’s range on the mainland. Lineage E included individuals from East China, both on the mainland and on islands in the East China Sea. Haplotypes from lineages W and E were co-distributed in Chuzhou and Chibi. The averaged pairwise distance of 6.23% between these lineages indicated a Miocene-Pliocene lineage-split. Lineage E was further subdivided to three sublineages: E1 and E2 comprised of haplotypes from the Zhoushan Islands, and E3 included haplotypes from the eastern mainland, the Zhoushan Islands and two islands south of the Zhoushan Islands. Lineages W and E showed evidence of demographic extensions. The isolation caused by the last transgression (0.01 Ma) has not yet led to a significant genetic differentiation between mainland and island populations in East China. However, divergence among some small islands may be driven by the restriction of migration and genetic drift.

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

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The lacertid lizard Takydromus wolteri widely distributed in South Korea, but intraspecific diversity of this species was not almost studied. T. wolteri represented by 8 specimens collected from 5 localities were analyzed by the Randomly Amplified Polymorphic DNA (RAPD) method with 28 random decamer primers and the phenogram constructed by using the Unweighted Pair Group Method with Arithmetic Mean (UPGMA) method based on RAPD data. The populations of T. wolteri using in this paper were collected from Gyeonggi-do, Chungcheongbuk-do, Jeju-do, Jeollanam-do and Gyeongsangnam-do in South Korea. Among 68 different bands detected in RAPD analysis, 59 bands (87%) showed polymorphism. The phenogram showed that the populations of T. wolteri were grouped into two. These results suggest that T. wolteri were supported intraspecific diversity.

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This study was conducted to find the sexual size dimorphism(SSD) in morphometric characters of three species , Eremias argus, Takydromus amurensis and T. wolteri in lacertid lizard from Korea. Six external traits, snout‐vent length, tail length, head length, head width, forelimb length and hind-limb length were focused to compare two sexes among the each three lizards. Student`s t‐test was used to compare the adult SVL between the sexes for each species. For the other parts of the body, a one‐way analysis of covariance(ANCOVA) with SVL as the covariate was used. In all these species, males have longer tail and head than females like general lacertid lizard. In addition, male E. arugus have broader head and longer forelimbs and hind-limbs than conspecific females. Likewise, male T. wolteri have broader head than females. In the three lacertid lizards from Korea, snout‐vent length did not significantly differ between the sexes. To understand the causes for SSD in SVL, ecological information, such as the presence/absence of male-male combat and the correlation between the SVL of female and litter size, is required. Therefore, further ecological study on the three species of lacertid lizard from Korea will make it possible to explain the reason SSD is not found in SVL.

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The present paper attempts to elucidate the taxonomic status of five species of Takydromus (T. amurensis, T. wolteri, T. tachydromoides oldi, T. kwagakunesis and T. auroralis) reported in Korean Peninsula to date. For this purpose 114 Takydromus specimens were collected from April 2003 to october 2004 in Southern Korean peninsula and morphological characters inclusive of the presence of contact between front-nasal and fostral and the number of femoral pores have been analyzed. The analysis reveals that T. kwagakunesis and T. auroralis should be synonymized to T. amurensis. In addition, the Korean Takydromus consists of two species of T. amurensis and T. wolteri.

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Anticipatory behaviour is the expectation of a near-future event based on information processed in the past and influences an animal’s tactical decisions, particularly when there are significant fitness consequences. The grass lizard (Takydromus viridipunctatus) perches on blades of grass at night which likely reduces the probability of predation by terrestrial preda- tors such as snakes, rodents and shrews. During twilight (starting 30 min before sunrise), they move from above the grass to within grass clumps and this is thought to afford the lizard protection while reducing detection by avian predators. Here, we examined how lizards shift their behaviour as a function of visual detectability to their primary predator, the cattle egret (Bubulcus ibis). We show that the lizards shift from their perch site during twilight at the earliest time at which egrets depart communal roosts. At the same time, visual modelling shows a dramatic increase in the detectability of the lizards to the visual system of egrets. Therefore, anticipatory behaviour in response to environmental cues acts to reduce pre- dation risk as lizards become more conspicuous and predators become more active. Grass lizard anticipatory behaviour appears to be finely tuned by natural selection to adjust to temporal changes in predation risk.

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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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Influence of body temperature on food assimilation and locomotor performance was studied in adult white-striped grass lizards (Takydromus wolteri) from a population in Chuzhou (Anhui, Eastern China). Food passage time dramatically decreased with increase in body temperature within the range from 26°C to 34°C, and then nearly levelled at higher body temperatures. Lizards overall gained net mass at 26°C, 28°C and 30°C and lost mass at 32°C and 34°C. Daily production of faeces (mass-specific) was affected by body temperature and, when influence of variation in the total food energy intake was removed by an ANCOVA, lizards at 26°C were found to produce faeces that contained significantly lower energy as compared with those at higher body temperatures. The total energy of urates was affected by the total food energy intake, but an ANCOVA with the total food energy intake as the covariate showed that lizards at different body temperatures did not produce urates that differed in energy. Within the temperature range considered, daily production of urates (mass-specific), apparent digestive coefficient (ADC) and assimilation efficiency (AE) were not affected by body temperature, although both ADC and AE were apparently greater in lizards at 26°C. Both locomotor stamina and sprint speed increased with increase in body temperature within the range from 18°C to 30°C, and then decreased at higher temperatures. Inter-individual differences were a significant source of variation in locomotor performance. Except for ADC and AE, all traits examined were significantly affected by body temperature, although thermal sensitivities differed considerably among traits. Within the body temperature range considered, 26°C was the most suitable body temperature for somatic tissue growth, and 30°C was an optimal body temperature for locomotor performance.

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The geographic distribution of species can be affected by environmental factors. The adult highaltitude lizard Takydromus hsuehshanensis (altitude > 1800 m) has been shown to tolerate summer lowland air temperature; its absence in lowland areas may therefore be caused by other factors. We employed a transplant experiment to monitor survival in different life stages and female reproduction in lowland areas. We maintained the T. hsuehshanensis adults in semi-natural outdoor enclosures with sufficient food, water, and the exclusion of potential predators. The results showed that (1) the survival rates of adults gradually decreased to 23.4% from one summer to the next, (2) illness occurred in adults during the winter, and (3) reproductive capacity (2 eggs/female), hatching success (31.7%), and hatchling survival rate (0% at the end of 11 weeks) were very low during the active season. We suggest that environmental factors synergistically caused these impairments in T. hsuehshanensis across different life stages and different seasons. This may partially explain its current altitudinal distribution.

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Among the life stages of oviparous reptiles, embryo and hatchling are particularly susceptible to temperatures. Since temperature decreases with altitude, the upper altitudinal limit of some lowland species could be related to the temperature requirement for embryogenesis. The endemic grass lizard Takydromus stejnegeri is widely distributed at altitudes below 1000 m in Taiwan. In this study, we examined the thermal effects on embryogenesis of this species and evaluated its potential influence on current altitudinal distribution of T. stejnegeri. We measured some indices of embryogenesis and hatchling after incubation at 5 constant temperatures (15, 20, 25, 30, and 33°C). The results revealed that (1) hatching success increased with temperatures from 15°C to 20°C, reached 100% at 25°C and 30°C, and then declined at 33°C; and (2) the incubation duration increased with decreasing temperatures. Based on polynomial regression analysis, we predicted the development of embryos would cease at 16°C. Comparing the monthly average temperature at 3 different altitudes, we found that the T. stejnegeri would have an incubation success <100 % at areas higher than 1000 m (monthly average soil temperature < 25°C), and its eggs would not have enough time to hatch in the areas above 1600 m. We assumed that the temperature requirement for embryogenesis could be a limiting factor for the current altitudinal distribution of T. stejnegeri.

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The present results indicate the name of Takydromus stejnegeri in Taiwan should be valid. With another close-related Taiwanese species (Takydromus formosanus), T. stejnegeri is distinct from Takydromus septentrionalis from mainland in the smaller body size but with the more longitudinal rows of dorsals. Takydromus stejnergeri is different from T. formonsanus in the lesser femoral pores and lesser longitudinal rows of dorsals extending to tail area.

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The female Takydromus stejnegeri in Taiwan has a definite breeding season from March to August . Most females produced two clutches but a few females may produce one or three clutches a year. The number of eggs per clutch for most clutches was 2 (with range from 1 to 4). The clutch size increased significantly during the breeding season. The smailest and largest mature females were 41mm and 57mm in snout-vent length, respectively. Most newly hatched individuals (<25mm in snout-vent length) occur from May to September. Fat body size was inversely correlated with reproduction. The mean liver weights peak in June. Both annual mean liver and fat body weights of T. stejnegeri females were similar to those of males during this same period.

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We describe a green lacertid lizard, Takydromus hani, from central Vietnam. It differs from congeners in having 6-8 (versus 1-5) femoral pores on each thigh. The phylogenetic relationships of this continental form seem to be with T. sauteri of Taiwan, and T. toyamai and T. smaragdinus of the Ryukyu Archipelago.

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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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Ein Bericht über die Terrarienhaltung und die Nachzucht von Takydromus smaragdinus BOULENGER, 1887.

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An inventory of amphibian and reptiles from the Barail Wildlife Sanctuary and its surroundings, Cachar District, Assam, north-eastern India is presented here. A total of 23 species of amphibians and 45 species of reptiles have been recorded from the study area. The observations include new locality records and natural history information of poorly known species such as Tropidophorus assamensis and Eutropis quadricarinata. Besides recording members of currently recognized species complex, the study also documents few species that were either conferred to closely related species (e.g., Calotes cf. irawadi) or their species identity remain to be ascertained (e.g., Rhabdophis sp. and Amolops sp.). The sharp slope in the species accumulation curve from the present study indicates that species count will rise further with additional surveys.

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In order to analysis and compare the effect of pregnancy and amputation of tail on temperature choice in the female ofTakydromus amurensis , the study was conducted in June and August of 2012. Two populations of Takydromus amurensis respectively from Shangzhi (Heilongjiang Province) and Kuandian (Liaoning Province) were studied by the method of field capturing and laboratory feeding. Our results showed that the Heilongjiang population ofT. amurensis could make their own body temperature increase through the heat source thermostat, when the temperature of the external environment was low (20℃ ). The mean temperature of the females were (32.49±0.705)℃ in the non-gestation period. However, in the pregnancy period, the mean temperature of the females were (33.90±0.674)℃, they were significantly higher after the pregnancy (F 1,61=65.600，P ＜0.001). No matter the pregnancy period or the non-gestation period, the mean temperature of the femalesT. amurensis from Shangzhi (33.39±0.909)℃ were significantly higher than the females from Kuandian (32.91±1.021)℃. It showed that the ability of thermoregulatory was stronger in the area of higher latitude. The mean temperature of the females from Kuandian were(31.94±0.590)℃, and that from Shangzhi were (31.60 ± 1.268)℃ after amputation of tail. The difference was not obvious between the two populations (F 1,18=0.591,P> 0.05), but the mean temperature was obviously lower than the individual which has the complete tail (F 1,18=9.854,P <0.05). The results showed that tail has influence on the thermoregulatory.

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Understanding the proximate determinants of phenotypic variations in life-history traits can provide powerful insights into a species’ life-history strategies. I experimentally manipulated availability of food (high vs low) to examine plasticity in the reproductive traits of northern grass lizards, Takydromus septentrionalis (Lacertidae), from eastern China. Food availability significantly affected reproductive frequency and thereby seasonal reproductive output, but had little effect on reproductive output per clutch. Low-food females postponed reproduction and produced less clutches in the reproductive season than did high-food females. After producing their second clutches, low-food females were in lower body condition than the high-food counterparts. By the end of the experiment, however, all females exhibited similar body condition. Clutch size and clutch mass differed between the first and second clutches but not between the treatments. Egg size and phenotypic traits of hatchlings (body size, morphology and locomotor performance) in T. septentrionalis did not vary significantly from first to second clutches nor between the two treatments. These results support optimal egg size (offspring) theory. Female T. septentrionalis “decide” whether or not to reproduce largely based on current energy intake; lowered feeding rates thus delay oviposition and reduce reproductive frequency. In contrast, clutch size, egg size and relative clutch mass remain unchanged.

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How fluctuating temperatures influence reptilian embryos and hatchlings has attracted increasing scientific attention, but is poorly known. We conducted an incubation experiment with a factorial design of two mean temperatures (24 vs. 28°C) and three diel thermal fluctuations (0, ±3, ±6°C) to determine the effects of diel thermal fluctuations and mean temperature on incubation duration and hatchling phenotypes. Both diel thermal fluctuations and mean temperature significantly affected incubation duration, but not hatching success. Incubation duration increased with increasing temperature fluctuations at a mean temperature of 24°C, but not at a mean temperature of 28°C. The significant interaction between diel thermal fluctuations and mean temperature on hatchling morphology indicated that the effect of thermal fluctuations depended on the mean temperature. Hatchling mass differed significantly between 24±6 and 28±6°C, but not between the two constant temperatures or the temperatures with ±3°C fluctuations. At a mean temperature of 24°C, the effect of thermal fluctuations on hatchling size was marginally significant, with relatively large hatchlings at the constant temperature; at a mean temperature of 28°C, thermal fluctuations had no impact on hatchling size. The locomotor performances were significantly affected by mean temperature rather than diel thermal fluctuations. Therefore, diel thermal fluctuations around a given mean temperature do not affect hatchling phenotypes in a relatively large magnitude, but the influence of diel thermal fluctuations may differ with changing mean temperatures.

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To understand how nest temperatures influence phenotypic traits of reptilian hatchlings, the effects of fluctuating temperature on hatchling traits must be known. Most investigations, however, have only considered the effects of constant temperatures. We incubated eggs of Takydromus septentrionalis (Lacertidae) at constant (24°C, 27°C, 30°C and 33°C) and fluctuating temperatures to determine the effects of these thermal regimes on incubation duration, hatching success and hatchling traits (morphology and locomotor performance). Hatching success at 24°C and 27°C was higher, and hatchlings derived from these two temperatures were larger and performed better than their counterparts from 30°C and 33°C. Eggs incubated at fluctuating temperatures exhibited surprisingly high hatching success and also produced large and well-performed hatchlings in spite of the extremely wide range of temperatures (11.6–36.2°C) they experienced. This means that exposure of eggs to adversely low or high temperatures for short periods does not increase embryonic mortality. The variance of fluctuating temperatures affected hatchling morphology and locomotor performance more evidently than did the mean of the temperatures in this case. The head size and sprint speed of the hatchlings increased with increasing variances of fluctuating temperatures. These results suggest that thermal variances significantly affect embryonic development and phenotypic traits of hatchling reptiles and are therefore ecologically meaningful.

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Geographic variation in offspring size is widespread, but the proximate causes of this variation have not yet been explicitly determined. We compared egg size and egg contents among five populations of a lizard (Takydromus septentrionalis, Günther, 1864) along a latitudinal gradient, and incubated eggs at two temperatures to determine the influence of maternal investment and incubation temperature on offspring size. The mean values for female size and egg size were both greater in the two northern populations (Chuzhou and Anji) than in the three southern populations (Lishui, Dongtou, and Ningde). The larger eggs were entirely attributable to the body size of females in the Anji population, but their increased size also stemmed from further enlargement of egg size relative to female body size in Chuzhou, the northernmost population sampled in this study. Eggs of the Chuzhou population contained more yolk and less water than those of southern populations. Despite the lower lipid content in the yolk, eggs from the Chuzhou population had higher energy contents than those from the two southern populations, owing to the larger egg size and increased volume of yolk. Hatchling size was not affected by incubation temperature, but differed significantly among populations, with hatchlings being larger in the Chuzhou population than in the other populations. Our data provide an inference that oviparous reptiles from cold climates may produce larger offspring, not only by increasing egg size but also by investing more energy into their eggs.

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Research on life-history traits of squamate reptiles has focused on North American species, while Asian taxa have been virtually ignored. In order to understand general patterns in reptile life histories, we need a broader data base. Our study on the slender-bodied lacertid lizard Takydromus septentrionalis provides the first detailed information on factors responsible for intraspecific variation in reproductive output and life history in a Chinese reptile. Clutches of recently collected lizards from five widely separated localities in China revealed major divergences in female body size at maturation, mean adult female body size, body condition after oviposition, size-adjusted fecundity, relative clutch mass, and mass and shape of eggs. Most of these geographical differences persisted when the same groups of females were maintained in identical conditions in captivity. Additionally, reproductive frequency during mainte- nance under laboratory conditions differed according to the animals’ place of origin. Thus, the extensive geographical variation in reproductive and life-history traits that occurs within T. septentrionalis is exhibited even in long-term captives, suggesting that proximate factors that vary among localities (local conditions of weather and food supply) are less important determinants of life-history variation than are intrinsic (presumably genetic) influences. The maternal abdominal volume available to hold the clutch may be one such factor, based on low levels of variation in Relative Clutch Mass among populations, and geographical variation in the position of trade-off lines linking off- spring size to fecundity.

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We used four species of lizards (Eumeces chinensis,Takydromus septentrionalis,Eremias brenchleyi and Calotes versicolor) that use different habitats as the experimental models to study variation in locomotor performance resulting from inter-specific differences in morphological traits.The sequence of body size measuring by snout-vent length (SVL) was E.chinensis (a ground-dwelling lizard)>C.versicolor (an arboreal lizard)>T.septentrionalis (a grass-dwelling lizard)>E.brenchleyi (a saxicolous lizard). Head size,limb length and tail length relative to SVL were the greatest in C.versicolor,intermediate in E.brenchleyi and T.septentrionalis,and the smallest in E.chinensis.Body height relative to SVL was the greatest in T.septentrionalis,intermediate in E.chinensis and C.versicolor,and the smallest in E.brenchleyi.On a horizontal racetrack,E.brenchleyi and T.septentrionalis run faster than did E.chinensis and C.versicolor.On an oblique racetrack,E.brenchleyi and C.versicolor run faster than did E.chinensis.The cling ability was the greatest in C.versicolor but the lowest in E.chinensis.Our data provide a support for the previous predictions that locomotion of animals has clear morphological determinants and that morphological variation may have fitness consequences through affecting locomotor performance.

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The body volume constraint hypothesis is a widely accepted notion that explains proximate limitation of female reproductive output, but lacks empirical support. To examine how body volume determines reproductive output, we experimentally manipulated the body volume of female lacertid lizard (Takydromus septentrionalis), to examine whether a reduction in body volume caused a reduction in reproductive output. Clutch size and mass decreased with a reduction in body volume. Reproductive output (residual clutch mass, RCM) was negatively related to body volume reduction (BVR) (RCM=−0.008BVR+0.225). In contrast, neither egg size nor offspring size was affected by the manipulation. Relative clutch mass was lower in manipulated females than in control females, but the actual physical burden (egg plus pearls) carried by manipulated females was much bigger than that of the control females. A normal clutch of eggs only filled 55% of maternal body volume, whereas manipulated females used 75% of body volume to contain eggs and pearls. Therefore, females are capable of carrying more eggs than a normal clutch. Our observations provide unequivocal empirical support for the maternal body volume constraint hypothesis. In addition, these results suggest that female T. septentrionalis may produce a clutch of eggs that fill the available abdomen space to an optimal level rather than the physical limit.

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Adult northern grass lizards (Takydromus septentrionalis) were collected at Mt.Xiaohe,Hangzhou in three periods of the breeding season,2001.They were dissected to determine sex,and storages of lipids in abdominal fat bodies,carcass,tail and liver.According to the stage of egg development (non-vitellogenetic follicles,yolked follicles,and oviductal eggs),females started vitellogenesis in later March (T1),produced eggs of the first clutch in early May (T2),and partly (37% of the sex) ceased reproduction in mid-June (T3).From T1 to T2,the abdominal fat body of females decreased from 17.3 mg on average to zero,and that of males changed from 6.3 mg to 0.8 mg.The decline of the lipid in females was 3 times of that of males.However,there was no significant change in lipid contents of the carcass,tail and liver in the three periods.It is very likely that the abdominal fat body of females is used for the first clutch of eggs,and earlier production of eggs means more clutches in the season and higher reproductive success.

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Understanding seasonal variation in reproductive investment is crucial to explore life-history strategies of a given species. We measured the timing of reproduction and the reproductive output of recently captured females to test whether the conclusions on seasonal shifts of reproductive investment made in the laboratory can apply to field populations in a lacertid lizard Takydromus septentrionalis. Female T. septentrionalis started producing eggs in April and ceased reproduction in August. During the midreproductive season (May to July), more than 60% of field-captured females were gravid. The females produced more eggs and thus had heavier total egg mass in May than in July, although egg mass and hatchling size did not vary seasonally. The seasonal variation in clutch size did not induce seasonal shifts in trade-offs between clutch size and egg size. In general, these results are largely consistent with data collected in the laboratory and support the conclusion of previous studies in this species that seasonal shifts in reproductive output are determined ultimately by natural selection. However, Relative Clutch Mass (RCM) of recently captured females was greater than that of females in captivity, which suggests that RCMs could be affected noticeably by proximate factors. Therefore, when explaining the evolutionary strategies of an organism based on intraspecific comparisons of RCMs, investigators should measure them in a commongardenenvironment so as to control the potential influence of environmental factors.

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We observed mating behavior of the Northern Grass Lizard(Takydromus septentrionalis) in an outdoor enclosure with aims to define the ethogram of mating behavior patterns and to determine the relationship between reproductive success and individual phenotypes in males.The general pattern of mating behavior in T.septentrionalis was:approaching → tail-biting → belly-biting → copulation.Duration of each behavior was 0.53 min,1.77 min,0.47 min and 141.3 min,respectively.Seventy percent of males that successfully mated were with greater mean snout-vent length and body mass than that of females.The mating success of males was positively correlated with their body size,but with their head size(length and width),tail length,body mass or skin color.

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Detecting inter-population differences in life-history traits is the first step in exploring the proximate and ultimate causes of such variation. We measured maternal body size and reproductive output of the lacertid lizard Takydromus septentrionalis from two island populations in eastern China to quantify inter-population variation. We captured female T. septentrionalis from the field and conducted a `common garden` experiment in the laboratory to measure their reproductive output. The study revealed major divergences in female body sizes, clutch mass and egg mass, but no significant difference in these traits was found between the first clutch and the later clutches. This suggests that the inter-population divergences persisted when the same groups of females were maintained in identical conditions in captivity. In contrast, there were no inter-population differences in size-adjusted fecundities, clutch size and relative clutch masses. Therefore, maternal body size plays an important role in determining female reproductive output in this species, but it does not account for all variation in reproductive traits. The egg size is less variable than the clutch size in each population, which gives support to the optimal egg size theory.

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The eggs of birds typically hatch after a fixed (but lineage-specific) cumulative number of heart beats since the initiation of incubation. Is the same true for non-avian reptiles, despite wide intraspecific variation in incubation period generated by variable nest temperatures? Non-invasive monitoring of embryo heart beat rates in one turtle species (Pelodiscus sinensis) and two lizards (Bassiana duperreyi and Takydromus septentrionalis) show that the total number of heart beats during embryogenesis is relatively constant over a wide range of warm incubation conditions. However, incubation at low temperatures increases the total number of heart beats required to complete embryogenesis, because the embryo spends much of its time at temperatures that require maintenance functions but that do not allow embryonic growth or differentiation. Thus, cool-incubated embryos allocate additional metabolic effort to maintenance costs. Under warm conditions, total number of heart beats thus predicts incubation period in non-avian reptiles as well as in birds (the total number of heart beats are also similar); however, under the colder nest conditions often experienced by non-avian reptiles, maintenance costs add significantly to total embryonic metabolic expenditure.

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We used Takydromus septentrionalis as a model animal to examine whether lizards shift thermal preferences when fasted. The experiment was conducted in March 2007. Thirty-six adults collected from Shengsi, Zhoushan Islands, were equally assigned to experimental (fasted) and control (fed) groups, with nine housed together in a 0.6 × 0.2 × 0.5 m3 glass cage with moist soil (10 cm in depth), debris and grasses. A 100-W light bulb suspended above one end of each cage created a thermal gradient ranging from room temperatures (15–24 °C) to 60 °C for 12 h daily, so that lizards could thermoregulate during the photophase. Prior to the experiment, lizards were allowed to feed on mealworms (larvae of Tenebrio molitor) for 10 days. Thereafter experimental lizards were fasted for 20 days, and were measured for thermal preferences on Day 1–10, Day 15 and Day 20 since fasting. We found in fasted lizards that selected body temperature (Tsel, a measure of thermal preference) gradually decreased from 32.4 °C to 30.9 °C in the first four days, and then increased to a level similar to that (32.9 °C) recorded in the controls. Lizards fasted for 9 days or longer did not differ from controls in Tsel. Our study is the first to demonstrate that lizards fasted for a prolonged time may shift thermal preferences in a pattern well fitting to a Gaussian peak curve

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From the publisher: Before you a unique atlas-determinant. For the first time in one edition descriptions of all species of amphibians and reptiles living on the territory of Russia are presented, illustrated with beautiful color photographs. The book is written by E.A. Dunaev and V.F. Orlova, the famous Russian herpetologists, researchers of the Zoological Museum of Moscow State University named after MV Lomonosov. Lomonosov. A simple and accessible form of presentation, detailed descriptions of the external appearance, lifestyle and behavior, photographs reflecting all variants of the variability of these animals, will help the curious reader to become more familiar with turtles, snakes, lizards, frogs and other representatives of the herpetofauna. And special keys will allow you to determine who you met on a country walk, in a city park, at a dacha or while traveling to different corners of our country. The book is of great interest not only for nature lovers, but also for specialists in herpetology and students of biological faculties of universities. It lists the amphibians and reptiles of the native fauna, reflects the latest changes in the taxonomy of these groups, lists species included in the Red Book of Russia and regional Red Books, lists reference and scientific publications on reptiles and amphibians of Russian fauna.

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A phylogeny of the family Lacertidae was derived from mtDNA gene sequence data. Seventeen species, representing 16 currently recognized genera and subgenera, were included in the analysis. A total of 954 bp was obtained and aligned from 12S and 16S partial gene sequences. A preferred tree was selected based on weighted parsimony and functional ingroup and outgroup analyses. Decay analysis, bootstrapping, and permutation tail probability were used to evaluate support for the recovered nodes. The genus Gallotia was resolved as the basal taxon and the sister group of all remaining lacertids.Takydromus branched off next. All African lacertids grouped together and formed a monophyletic clade with the Eurasian genera Eremiasand Ophisops. The remaining Eurasian lacertids sequentially branched off near the base of the tree in a “comb-like” fashion. The basal position of Gallotia and the monophyly of African lacertids are consistent with previous hypotheses. The European-origin hypothesis of lacertids is favored, and the distribution of lacertids in Africa is likely a Miocene dispersal event. Most of the extant European lacertids probably arose after the Eocene. The classification of the family needs to be revised.

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The feeding and development of Amblyomma testudinarium Koch larvae on the lizard, Takydromus tachydromoides (Schlegel) were observed at 24℃ under a 16L-8D photoperiod and compared with those of the larvae on mice. The mean feeding periods of the larvae were 13.7 and 5.4 days on the lizards and the mice, respectively. The mean feeding period on the lizards was significantly longer than that on the mice. However, the molting success of the engorged larvae was 97% or more on both hosts. The mean developmental periods of the engorged larvae were 22.4 and 22.6 days on the lizards and the mice, respectively. The difference in the mean developmental period between the two hosts was not significant, suggesting that the difference in the host suitability between the two hosts is slight. Thus, the reasons why the immature ticks of A. testudinarium infest more frequently on mammals than on reptiles can not be explained by the difference in the host suitability between the cold-blooded and warm-blooded animals.

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The Japanese lacertid lizard Takydromus tachydromoides and the praying mantis Tenodera aridifolia are sympatric generalist predators feeding on similar prey. To confirm reciprocal predation between them, we observed the behavioural interactions between the lizards and the mantises of different sizes in a laboratory condition. The lizards caught small mantises (from first to fifth instars), but sometimes escaped from large mantises (from sixth instar to adult). Large mantises occasionally showed catch responses to the lizards. The lizards sometimes caught the mantis without a tongue-flick response (sampling of chemical cues), and they sometimes did not catch the small mantises showing immobile or cryptic responses that prevent visual detection. These results suggested the primary role of vision on recognition of the mantis as a prey. The lizards spent a longer time to approach larger mantises. The time from orienting to catch was longer when the lizards showed tongue-flick responses. The lizard also spent a longer time before deciding to escape from the mantis than to catch it. Biological significance of these differences in timing was discussed.

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It is important to investigate visual orienting in reptiles to better understand the basic organization of the oculomotor system in vertebrates. However, quantitative analyses of visual orienting behavior in reptiles have rarely been conducted, except in chameleons. In the present study, we videorecorded the head and body movements of the lizard Takydromus tachydromoides during visual tracking of moving prey and analyzed them frame-by-frame. Before approaching prey, visual tracking mainly consisted of brief intermittent turns of the head (saccade). After the head saccades, the angular position of the prey relative to the lizard head was kept at 10–70° (laterally) in most cases, rather than at 0° (in front). In addition, the ratio of the amplitude of the head saccades to prey position was 0.2–0.3, which is much smaller than 1, suggesting that the head did not orient exactly toward the prey after most saccades. These results were observed under both white (homogeneous) and grating (structured) backgrounds. Possible functions of head saccades in the lizard are discussed.

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

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We report the first record of Sikkim grass lizard, Takydromus sikkimensis Günther, 1888 in Nepal based on morphological characters such as the presence of four pairs of femoral pores, 12 rows of ventral scales, tail more than 3.6 times longer than snout-vent length. Our record of T. sikkimensis at Miklajung, Morang district represents the western-most observation of the species, ca. 94 km west of its type locality, Sikkim, India and is the first in Nepal for this species, genus, and family. This record is from Chure/Siwalik hill range which lies outside of Nepal’s protected area network. This species is listed as Endangered by the International Union for Conservation of Nature and Natural Resources and warrants detailed inventory and immediate conservation interventions.

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

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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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Background: Identifying the factors that contribute to divergence among populations in mate preferences is important for understanding of the manner in which premating reproductive isolation might arise and how this isolation may in turn contribute to the evolutionary process of population divergence. Here, we offered female northern grass lizards (Takydromus septentrionalis) a choice of males between their own population and another four populations to test whether the preferences that females display in the mating trials correlate with phenotypic adaptation to local environments, or to the neutral genetic distance measured by divergence of mitochondrial DANN sequence loci. Results: Females showed a strong preference for native over foreign males. Females that mated with native versus foreign males did not differ from each other in mating latency, or copulation duration. From results of the structural equation modelling we knew that: 1) geographical distance directly contributed to genetic differentiation and environmental dissimilarity; 2) genetic differentiation and environmental dissimilarity indirectly contributed to female mate preference, largely through their effects on morphological divergence; and 3) females judged mates by body shape (appearance) and discriminated more strongly against morphologically less familiar allopatric males. Conclusions: Local adaptation rather than neutral genetic distance influences female mate preference in T. Septentrionali. The tendency to avoid mating with foreign males may indicate that, in T. septentrionalis, local adaptations are more valuable than genetic novelties. Our results highlight the importance of comprehensive studies integrating ecological, molecular and behavioral approaches to understand population divergence in female mate preferences as the consequence of local adaptations.

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Background: Identifying the factors that contribute to divergence among populations in mate preferencesis important for understanding of the manner in which premating reproductive isolation might arise andhow this isolation may in turn contribute to the evolutionary process of population divergence. Here, weoffered female northern grass lizards (Takydromus septentrionalis) a choice of males between their ownpopulation and another four populations to test whether the preferences that females display in themating trials correlate with phenotypic adaptation to local environments, or to the neutral geneticdistance measured by divergence of mitochondrial DNA sequence loci. Results: Females showed a strong preference for native over foreign males. Females that mated withnative versus foreign males did not differ from each other in mating latency, or copulation duration. Fromresults of the structural equation modelling we knew that: 1) geographical distance directly contributed togenetic differentiation and environmental dissimilarity; 2) genetic differentiation and environmentaldissimilarity indirectly contributed to female mate preference, largely through their effects onmorphological divergence; and 3) females judged mates by body shape (appearance) and discriminatedmore strongly against morphologically less familiar allopatric males. Conclusions: Local adaptation rather than neutral genetic distance influences female mate preference in T. septentrionalis. The tendency to avoid mating with foreign males may indicate that, in T.septentrionalis, local adaptations are more valuable than genetic novelties. Our results highlight theimportance of comprehensive studies integrating ecological, molecular and behavioral approaches tounderstand population divergence in female mate preferences as the consequence of local adaptations.

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The northern grass lizard (Takydromus septentrionalis) is an oviparous lacertid lizard that is endemic to China and has a wide distributional range covering the central and south-eastern parts of the country. It is listed on the ‘‘lists of terrestrial wildlife under state protection, which are beneficial or of important economic or scientific value’’ promulgated by China’s State Forestry Administration. Here, nine polymorphic microsatellite loci were isolated from T. septentrionalis genomic libraries. These markers revealed a high degree of genetic diversity (2–22 alleles per locus) and heterozygosity (H O ranged from 0.063 to 0.938, and H E ranged from 0.06 to 0.928). No locus exhibited deviations from Hardy–Weinberg equilibrium. There was no evidence of linkage disequilibrium among pairs of loci. These microsatellite markers will be useful for future study of population genetic analysis and parentage assignment of T. septentrionalis.

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During our collecting trip to Guangxi in 2016, we collected ten specimens of the genus Takydromus from the suburb of Guilin, northeastern Guangxi, South China, and found that they did not belong to any currently known species. Here, we described this new species, Takydromus guilinensis sp. nov., based on morphological and mitochondrial DNA (CO1 and cyt b) data. This new species is a sister taxon to T. intermedius with a p-distance of 0.070 in CO1 and 0.080 in cyt b. These two p-distances exceed not only the minimum value (0.067) between /i>T. septentrionalis and T. stejnegeri but also the minimum value (0.079) between T. intermedius and T. yunkaiensis. Morphologically, this new species differs from other currently recognized Takydromus species from the same clade, more evidently in the longitudinal rows of dorsal scales, transverse rows of scales at the mid-body and mensural variables. The description of Takydromus guilinensis sp. nov. increases the total number of Takydromus species to 25, of which 16 can be found in China. Takydromus guilinensis sp. nov. is currently known only from Guilin, Guangxi, South China, where it is sympatric with the other four Takydromus species (T. septentrionalis, T. kuehnei, T. sexlineatus and T. intermedius).

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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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One of the main functions of physiological color change is thermoregulation.This change occurs much more rapidly than morphological color change, but the underlying mechanism remains poorly understood. Here, we studied the thermal dependence and molecular basis of physiological color change inlizards using Takydromus septentrionalis (Lacertidae) as the model system. Body color was thermally sensitive, becoming increasingly light as body temperatures deviated from the level (~30 °C) preferred by this species. We identified 3,389 differentially expressed genes (DEGs) between lizards at 24 °C and 30 °C, and 1,097 DEGs between lizards at 36 °C and 30 °C. Temperature affected the cAMP signal pathway, motor proteins, cytoskeleton, and the expression of genes related to melanocyte-stimulating hormone (MSH) and melanocyte-concentrating hormone (MCH). Our data suggest that the role of physiological color change in thermoregulation is achieved in T. septentrionalisby altering the arrangement of pigments and thus the amount of solar radiation absorbed and reflected. G protein-coupling system inhibits adenylate cyclase activity to transform ATP into cAMP and thereby causes rapid pigment aggregation. MCH deactivates the G proteins and thereby initiates pigment dispersion. This mechanism differs from that reported for teleost fish where MCH activates the G proteins and thereby causes pigment aggregation.

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Thermal biology traits reflect thermal adaptations to an environment and can be used to infer responses to climate warming in animal species. Within a widespread genus or species, assessing the latitudinal or altitudinal gradient of thermal physiological tra its is essential to reveal thermal adaptations and determine future vulnerability to climate warming geographically. We determined the thermal biology traits of a cold-climate distributed lizard, Takydromus amurensis, and integrated published thermal biology traits within the genus Takydromus to reveal a preliminary geographical pattern in thermal adaptation. The mean selected body temperature (cloaca temperature; Tsel), critical thermal maximum (CTmax), critical thermal minimum (CTmin), and optimal temperature for locomotion (i.e., sprint speed; Topt) of T. amurensis were 32.6, 45.1, 3.1, and 33.4 °C, respectively. The resting metabolic rates of T. amurensis were positively related to temperature from 18 °C to 38 °C. We compared the traits of tropical T. sexlineatus, subtropical T. septentrionalis, and T. wolteri with T. amurensis and found that the CTmax and thermal tolerance range (the difference between CTmax and CTmin; TTR) increased toward high latitudes, whereas CTmin increased toward low latitudes in these four Takydromus lizards. According to this preliminary pattern, we speculate the species at medium and low latitudes would be more vulnerable to extreme heat events caused by ongoing climate warming. We highlight the importance of integrating thermal biology traits along geographical clues, and ist potential contribution to evaluate the vulnerabilities of species in the context of climate warming.

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

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

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

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The complete mitochondrial genome sequence of Takydromus septentrionalis was determined by shotgun sequencing. The total length of mitogenome is 18,304 bp, and contains 13 protein-coding genes, 22 tRNA genes, 2 ribosome RNA genes, and 2 control regions. Most of the genes of T. septentrionalis were distributed on the H-strand, except for the ND6 subunit gene and eight tRNA genes which were encoded on the L-strand. The phylogenetic tree of T. septentrionalis and 8 other closely related species was reconstructed. The phylogenetic analyses based on these mitogenomes presented here will be useful for further insights on the evolutionary relationships of Takydromus.

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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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1. We investigated whether heat tolerance has a crucial impact on the altitudinal distribution of a high-altitude lizard, Takydromus hsuehshanensis (>1800 m altitude). 2. We measured and compared its heat tolerance with that of another two lowland species. Heat tolerance measurement included critical thermal maximum (CTMax) and survival rates under three fluctuating daily temperature treatments over a 3-month period. Two of the three temperature treatments, the extremely high temperature (EH treatment) and the average temperature (H treatment), were set to approximate lowland summer temperatures. The third one was set to approximate cool temperature in mountain areas (C treatment). 3. Our results showed that (1) CTMax of T. hsuehshanensis was higher than the summer temperature in the lowland areas, and not significantly lower than that of two other lowland species, (2) T. hsuehshanensis survived the H and EH treatments over a 3-month period and its survival rate was not significantly lower than that of the other two lowland species. Therefore, T. hsuehshanensis was not only able to tolerate high temperatures mimicking lowland areas for a short period of time, but also for a much longer period of time. 4. We conclude that the heat tolerance of T. hsuehshanensis is not a crucial factor limiting its current altitudinal distribution.

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We investigated whether cold tolerance was an important limiting factor in the current altitudinal distributions of two Takydromus lizards, T. formosanus (< 1500 m in altitude) and the lowland-dwelling T. stejnegeri (< 1000 m in altitude) in Taiwan. We measured their critical thermal minimum (CTMin) and 3 mo survival rates in 4 cold treatments, and compared these with a high-mountainous species, T. hsuehshanensis (> 1800 m in altitude). The results indicated that (1) both the CTMin and prolonged cold tolerance were correlated to their upper limit of altitudinal distributions as predicted and (2) T. formosanus and T. stejnegeri had reasonable survival rates at temperatures that were lower than the underground temperature of high altitudinal areas. We concluded that although cold tolerance was correlated with altitudinal distribution, it is not a crucial factor limiting T. formosanus and T. stejnegeri at higher altitudes.

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We investigated the impact of environmental temperature on elevational distributions of a high-mountainous lizard, Takydromus hsuehshanensis, by examining the thermal sensitivity of its locomotor performance. Its sprint speed was measured at 9 body temperatures after 2 wk of acclimation at 2 different temperatures. The same measurements were performed on a closely related species, T. formosanus, which lives at lower elevations. The results indicated that (1) T. hsuehshanensis was capable of maintaining normal locomotor performance within a body temperature range which approximates summer temperatures of lowland areas; (2) T. formosanus was able to run significantly faster than T. hsuehshanensis within a certain range of body temperatures; (3) these 2 species did not differ in their thermal sensitivities of the locomotor performance; and (4) temperature acclimation treatments did not affect the locomotor performance of these 2 species. We concluded that locomotor performance is not a crucial factor limiting the distribution of T. hsuehshanensis in lowland areas.

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Warmer climates have affected animal distribution ranges, but how they may interact with vegetation patterns to affect habitat use, an important consideration for future wildlife management, has received little attention. Here, we use a biophysical model to investigate the potential thermal impact of vegetation pattern on the habitat quality of a high-elevation grassland lizard, Takydromus hsuehshanensis, and to predict the thermal suitability of vegetation for this species in a future warmer climate (assuming 3 °C air temperature increase). We assess the thermal quality of vegetation types in our study area (Taroko National Park in areas >1,800 m) using three ecologically relevant estimates of reptiles: body temperature (T b), maximum active time, and maximum digestive time. The results show that increasing forest canopy gradually cools the microclimates, hence decreasing these estimates. In the current landscape, sunny mountain-top grasslands are predicted to serve as high quality thermal habitat, whereas the dense forests that are dominant as a result of forest protection are too cold to provide suitable habitat. In simulated warmer climates, the thermal quality of dense forests increases slightly but remains inferior to that of grasslands. We note that the impact of warmer climates on this reptile will be greatly affected by future vegetation patterns, and we suggest that the current trend of upslope forest movement found in many other mountain systems could cause disadvantages to some heliothermic lizard species.

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In this study, I describe seasonal variations in the male and female reproductive cycles of Takydromus hsuehshanensis, an oviparous grass lizard endemic to the high altitude areas of Taiwan. Mean snout-vent length (SVL) of adult males (n = 78) and females (n = 105) were 60.5 mm (range = 47-72 mm) and 61.2 mm (range = 54-72 mm), respectively. Females exhibited a spring vitellogenic period with parturition occurring in May to July. The onset of vitellogenesis showed a negative correlation with the mass of the female fat body. Females produced two to four eggs per clutch, and clutch size was positively correlated with SVL: only females with SVLs equal to or larger than 61 mm produced a clutch consisting of four eggs. Two clutches were recorded during a single year in some individuals. Males exhibited a continuous spermatogenic pattern, but testis mass, accessory sexual organ mass, and seminiferous tubule diameter showed cyclical changes with a peak before the dormant months from September to December. The tubules of the epididymides were full of sperm from December to the following March, indicating that sperm were stored in the epididymides until the occurrence of ovulation in the spring. Male fat bodies exhibited the lowest mass from March to June, coincident with the period of reproductive activity. Male seminiferous tubule diameter exhibited a positive correlation with liver, fat body, and testis masses. However, seminiferous tubule diameter, as well as testes mass, had a negative correlation with precipitation. Timing of reproduction in four other sympatric lizards and clutch size variation in other Takydromus species are compared with those in T. hsuehshanensis.

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I describe habitat use, diet, and male and female reproductive cycles of Takydromus sauteri, an oviparous grass lizard inhabiting Orchid Island, a tropical island off the southeastern coast of Taiwan. All T. sauteri were observed on leaves of Japanese silver-grass (Miscanthus floridulus) at the forest edge. The diet of T. sauteri consisted mostly of spiders (35.7%) and Hemipterans (14.3%). Two prey categories, spiders and gymnoceratans, numerically dominated the diet. Two lizards` stomach contained plant seeds, and two had eaten roaches. Mean snout-vent length (SVL) of adult males (N = 59) was 64.2 (range: 60.3-73.7) mm and that of females (N = 63) was 67.1 (range: 50.6-76.5) mm. Females exhibited spring and summer vitellogenesis with parturition occurring from February to August. Onset of vitellogenesis showed a negative correlation with mass of female fat bodies. Females produced one to four eggs per clutch, and clutch size was not correlated with SVL. Two clutches were recorded during a single year in some individuals. Male fat bodies exhibited the lowest mass from January to July, coincident with the period of reproductive activity.

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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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Scanning electron microscopy was employed to investigate the ultrastructure of oral epithelialcells of the lizard, Takydromus tachydromoides. The specimens were prepared by a methodinvolving osmium postfixation and acid treatment to remove extracellular attached material. Bump-shaped protuberances were arranged symmetrically on both sides of the median line of thepalate. Epithelial surfaces besides the bump-shaped protuberances were widely covered with fine folds.At higher magnification, a network pattern of microridges was closely distributed over the entiresurface of the palate, and each cell marginal thickening was clear. The protuberances were alsoobserved in the latero-posterior region of the floor of the mouth. A number of nicks were arrangedsymmetrically on both sides along the median line of the fore-region of the floor. Taste buds werescattered in the epithelial surfaces between nicks. At higher magnification, microvilli as well as microridgeswere observed on the epithelial cell surfaces of the floor. There were no papillar structures onthe ventral tongue surface. Fine pits were densely distributed over the entire epithelial cellsurface. The cell margin was clearly distinguishable as a depressed line. The epithelial surface ofthe laryngeal part of the pharynx was entirely covered with ciliated cells. Taste buds were alsoscattered on the inner gingival epithelial surfaces of the upper and lower jaws along the dentalarches. Relatively indistinct microridges were widely distributed on the gingival epithelial cell surfaceof the upper jaw. Scrollwork and network patterns of microridges were clearly observed on the gin -gival epithelial surface of the lower jaw.

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The Japanese lacertid Takydromus tachydromoides is an opportunistic, generalized predator; its diet consists chiefly of spiders and insects although other small arthropods and gastropods may be taken. Prey are lognormally distributed by size. Age-, season- and habitat-specific dietary differences are intimately related to the habits of this species. Such differences occur primarily in response to prey densities and are of adaptive value to the lizards. Most prominent among these differences is the ratio of insects to spiders in the diet. Competition between adult and juvenile lizards is reduced through partial nonoverlap of food resources and structural habitats. Takydromus displays certain aspects of predation recognized in the past predominantly from larger and better-known mammalian and reptilian carnivores. As a predator Takydromus probably exerts little limiting influence on its prey.

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Rare or newly discovered species are attractive targets for poaching as they often command a high value in the international market. Endemic species often have a very restricted range, making them extremely vulnerable to stochastic events and human activities, including poaching. Previous studies have shown that several species endemic to the Nansei Islands in Japan are available in the international pet trade. This paper further investigates the presence of live reptiles and amphibians endemic to the Nansei Islands in international pet trade. Seven Facebook groups and five classified websites were searched for these species, in addition to the CITES database and a spot-check at Chatuchak market in Thailand. Nansei Island endemic reptiles and amphibians are more common in the international pet trade than initially thought. During the study period, advertisements for 23 species endemic to the Nansei Islands were found, comprising a total of 360 specimens. In addition, during a spot-check, 8 individuals of 2 subspecies of Japanese sword-tail newt were found at Chatuchak market in Thailand. Only three of these species are protected by the CITES and nine are regionally or nationally protected. Yet, ten species observed in international markets are classified as Endangered by IUCN and two even as Critically Endangered. The large numbers observed for the Endangered and Critically Endangered Goniurosaurus and Takydromus species warrants increased protection and a CITES Appendix III listing is recommended for these species.

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Recently, amphibians and reptiles have drawn attention because of declines in species and populations caused mainly by habitat loss, overexploitation and climate change. This study constructed a DNA barcode database for the Korean herpetofauna, including all the recorded amphibians and 68% of the recorded reptiles, to provide a useful, standardized tool for species identification in monitoring and management. A total of 103 individuals from 18 amphibian and 17 reptile species were used to generate barcode sequences using partial sequences of the mitochondrial cytochrome c oxidase subunit I (COI) gene and to compare it with other suggested barcode loci. Comparing 16S rRNA, cytochrome b (Cytb) and COI for amphibians and 12S rRNA, Cytb and COI for reptiles, our results revealed that COI is better than the other markers in terms of a high level of sequence variation without length variation and moderate amplification success. Although the COI marker had no clear barcoding gap because of the high level of intraspecific variation, all of the analysed individuals from the same species clustered together in a neighbour-joining tree. High intraspecific variation suggests the possibility of cryptic species. Finally, using this database, confiscated snakes were identified as Elaphe schrenckii, designated as endangered in Korea and a food contaminant was identified as the lizard Takydromus amurensis.

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1. During incubation, rigid-shelled eggs of Gekko japonicus declined in both wet mass and dry mass, and pliable-shelled eggs of Takydromus septentrionalis increased in wet mass but decreased in dry mass. 2. Rapid increase of percentage of lipids in developing eggs occurred during the early period of vitellogenesis, percentage of lipids in medium and large sized follicles of both species remained approximately constant. Caloric value was correlated with lipid content and for ash, uncorrected caloric value was correlated with ash content. 3. Total ash present in developing eggs increased as the eggs developed. Embryos of both species did not rely exclusively on ash supplied in their yolks, and withdrew ash from their eggshelis.

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An experimental reduction of offspring number has been reported to result in enlargement of offspring size in lizards. We applied the ‘‘follicle excision’’ tech- nique to a lacertid lizard (Takydromus septentrionalis) to examine whether this effect is generalisable to lizards. Of the 82 females that produced 3 successive clutches in the laboratory, 23 females underwent follicle excision after they oviposited the first clutch. Follicle excision reduced clutch size, but did not alter egg size. This result indi- cates that egg size is not altered during vitellogenesis in T. septentrionalis. Females undergoing follicle excision produced a third clutch (a second post-surgical clutch) as normally as did control females. Females switched from producing more but smaller eggs early in the breeding season to fewer but larger eggs later in the season. Our results indicate that female T. septentrionalis maximize reproductive success by diverting an optimal, rather than a higher, fraction of the available energy to individual offspring. This optimized allocation of the available energy to offspring production explains why follicle excision does not result in enlargement of egg size in this species. Our study provides evidence that an experimental reduction of offspring number does not always result in enlargement of offspring size in lizards.

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We measured the reproductive output of Takydromus septentrionalis collected over 5 years between 1997 and 2005to test the hypothesis that reproductive females should allocate an optimal fraction of accessible resources in a par-ticular clutch and to individual eggs. Females laid 1–7 clutches per breeding season, with large females producingmore, as well as larger clutches, than did small females. Clutch size, clutch mass, annual fecundity, and annualreproductive output were all positively related to female size (snout–vent length). Females switched from producingmore, but smaller eggs in the first clutch to fewer, but larger eggs in the subsequent clutches. The mass-specific clutchmass was greater in the first clutch than in the subsequent clutches, but it did not differ among the subsequentclutches. Post-oviposition body mass, clutch size, and egg size showed differing degrees of annual variation, butclutch mass of either the first or the second clutch remained unchanged across the sampling years. The regressionline describing the size–number trade-off was higher in the subsequent clutch than in the first clutch, but neither theline for first clutch, nor the line for the second clutch varied among years. Reproduction retarded growth more mark-edly in small females than in large ones. Our data show that: (1) trade-offs between size and number of eggs andbetween reproduction and growth (and thus, future reproduction) are evident in T. septentrionalis; (2) females allo-cate an optimal fraction of accessible resources in current reproduction and to individual eggs; and (3) seasonal shiftsin reproductive output and egg size are determined ultimately by natural selection.

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1. We studied in field and laboratory experiments several aspects of the thermal biology of adult lacertid lizards (Takydromus septentrionalis). 2. The selected body temperature and the critical thermal maximum averaged 30.0 and 42.3°C, respectively; the critical thermal minimum of males and females averaged 3.9 and 5.9°C, respectively. 3. The lizards in May exhibited lower and more variable active body temperatures than did the lizards in July, because of lower and more variable air and substrate temperatures in May. 4. The sprint speed generally increased with body temperature within the range of 17 to 32°C and decreased at temperatures >32°C. 5. Within the range of 26 to 36°C, body temperature influenced food assimilation primarily by affecting the food passage time, whereas it only slightly affected the daily food consumption, the apparent digestive coefficient and the assimilation efficiency. The food passage time decreased with increasing body temperature.

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Home range sizes of the lizard Takydromus Septentrionalis were evaluatedduring reproduction period in 1987 and 1988 at Xiushan Island in Zhejiang province. Most of the lizard occupied a certain area of home range. Mean squared recapture radii (r2) were used to estimate home range size. In 1988, adults had greater home range sizes than juveniles, the mean value of r2 of the latter being 11.6±7.3SD. Adult males had greater home range sizes than adult females, the mean values being 49.8±13.9SD and 31.1±22.2SD, respectively. In 1987,adult males also had greater home range than adult females, the mean values of r2 being 66.1±16.7SD and 35. 2±19. 1SD, respectively. Population size in 1987 was greater than that in 1988,there was an inverse relationship between population size and home range size,although there was no significant difference in mean recapture radii both in males and in females.

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1. We studied in field and laboratory experiments several aspects of the thermal biology of adult lacertid lizards (Takydromus septentrionalis). 2. The selected body temperature and the critical thermal maximum averaged 30.0 and 42.3°C, respectively; the critical thermal minimum of males and females averaged 3.9 and 5.9°C, respectively. 3. The lizards in May exhibited lower and more variable active body temperatures than did the lizards in July, because of lower and more variable air and substrate temperatures in May. 4. The sprint speed generally increased with body temperature within the range of 17 to 32°C and decreased at temperatures >32°C. 5. Within the range of 26 to 36°C, body temperature influenced food assimilation primarily by affecting the food passage time, whereas it only slightly affected the daily food consumption, the apparent digestive coefficient and the assimilation efficiency. The food passage time decreased with increasing body temperature.

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1. Juvenile Takydromus septentrionalis at 28, 30 and 32°C exhibited no significant differences in food intake but significant differences in weight-specific food intake, with the highest food intake (384.6 cal/day) and the highest weight-specific food intake (195.2 cal/g/day) occurring at 32 and at 30°C, respectively. 2. The assimilation efficiencies at 28, 30 and 32°C averaged 87.7, 88.7 and 83.3%, respectively. 3. No significant differences in growth were observed among lizards at different temperatures.

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Sexual dimorphism and reproduction in a population of Takydromus septentrionalis on Xiushan Island of Zhoushan Islands group were investigated during the breeding seasons of 1989 - 1992. Most males and females, approximately two years after hatchling, reached sexual maturity at 54 and 57 mm SVL, respectively. Adult male and female T. septentrionalis were similar in SVL but quite different in the head size characteristics, males having larger heads than females of similar SVL except near the size of sexual maturity. Mating success in males was apparently not related to increased SVL. Males can copulate with females in several hours before or after oviposition. The duration of intromission averaged 227 min. Females can lay 1 - 3 clutches with 2 - 6 eggs each per season. Clutch size or clutch mass was significantly correlated with SVL and was not correlated with the clutch size. Relative clutch weight (RCW) averaged 0.149. Oviposition frequency averaged 16.7 days.

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The prey recognition system in the predatory behavior of T. tachydromoides was analyzed experimentally using unpalatabe Pryeria moths and edible mealworms. None of the Pryeria moths were either killed or eaten, while all the mealworms attacked were eaten. The learning of visual avoidance by T. tachydromoides, in which the lizard associates the unpalatability of Pryeria moth with its appearance and avoids it, seemed to exist, but its process appeared to be very slow. In the course of predatory behavior, the lizard first recognized the prey visually, but then checked it strictly through chemical cues. This suggested that not only visual but also other sensory cues such as olfactory and gustatory stimuli derived from the prey were necessary for the efficient avoidance learning in this lizard.

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

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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 investigated sexual size dimorphism (SSD), sexual shape dimorphism (SShD), and allometric growth in the skull of Takydromus tachydromoides using geometric morphometries. We analyzed 40 landmarks on the skull from dorsal, ventral, and lateral views. As a result, significant sexual dimorphisms were recognized in both size and shape of the skull. The analysis of SSD revealed that adult males have larger skull relative to the snout-vent length, and the analysis of SShD indicated that adult males have larger jaw muscle adductor chambers than adult females. Moreover, various allometric shape changes in both sexes were visualized by the thin plate splines. However, only subtle sexual differences in allometric shape changes were observed. The skull size and shape of this species may be influenced by phylogenetic constraints and only weakly driven by sexual selection.

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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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To identify the biological characteristics of Wolter lizard (Takydromus wolteri), this study was conducted from March 2011 to August 2012 in the areas of Geumoreum located in Hangyeong-myeon, and Saebyeoloreum located in Aewol-eup and in the zoological laboratory in Jeju National University, South Korea. Food source, enemies, activity period, breeding period, spawning period, spawning size in a clutch, biological traits of eggs and hatching periods were investigated. As a result of the study, it was found that T. wolteri eats Gryllidae, Araneina, Orthoptera, Diptera, Hymenoptera and Lepidoptera, but not Coleoptera, Formicidae, Oligochaeta. By observing the vicinities of T. wolteri habitats, it was seen that the natural enemies of T. wolteri were Elaphe dione, Amphiesma vibakari ruthveni, Sibynophis chinensis, Rhabdophis tigrinus, Gloydius ussuriensis, Corvus macrorhynchos, Streptopelia orientalis, Falco tinnunculus, Pica pica sericea, Lanius bucephalus, Hypsipetes amaurotis. Activities started in mid-March and hibernation started between late November and early December. Mating period was seen to run between May to June, breeding was May to August, and spawning was June toAugust. The average weight of spawned eggs (n = 12) was 0.3g; size was 9.5 × 7.2 mm; and the average fecundity of the clutch was 5 (with range 3~7). Incubation period was 18~20 days. The results would be useful as the necessary material for preparing conservation and management plans for T. wolteri habitats that depend on the changes in the habitat environment.

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

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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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Recent studies have demonstrated that thermoregulatory behavior occurs not only in posthatching turtles but also in turtles prior to hatching. Does thermoregulatory behavior also occur in the embryos of other reptile and bird species? Our experiments show that such behavior is widespread but not universal in reptile and bird embryos. We recorded repositioning within the egg, in response to thermal gradients, in the embryos of three species of snakes (Xenochrophis piscator, Elaphe bimaculata, and Zaocys dhumnades), two turtles (Chelydra serpentina and Ocadia sinensis), one crocodile (Alligator sinensis), and four birds (Coturnix coturnix, Gallus gallus domesticus, Columba livia domestica, and Anas platyrhynchos domestica). However, we detected no significant thermoregulation by the embryos of two lizard species (Takydromus septentrionalis and Phrynocephalus frontalis). Overall, embryonic thermoregulatory behavior is widespread in reptile as well as bird species but may be unimportant in the small eggs laid by most lizards.

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The relationship between form, function and ecology is very important for us to understand the process of phenotypic evolution and ecological evolution in biological organisms. Trait evolution between species and populations is the result of multiple factors. The Bite force of a vertebrate plays an important role in its life, mainly including eating habits, mate competition, territorial possession, defense against invaders and so on. In this study, the northern grass lizard (Takydromus septentrionalis) was used as a subject to explore the influence of living environmental factors on lizard morphology and bite force. After measuring the morphological traits of five populations, it was made a comparative analysis, on the differences in morphology and bite force between populations by studying the morphology, bite force and sexual dimorphism between different populations. The results showed that with the morphological size and bite force of the lizard increase as the average annual temperature of the land population decreases, however, the island population did not show this phenomenon. A significant positive correlation between the head size and the bite force of the lizard was found,and the obvious sexual dimorphism of head size,mouth opening and bite force of the lizard was also detected.

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A female grass lizard Takydromus stejnegeri van Denburgh was captured after mating in the field on 24 April 2003. It was immediately brought back to the laboratory and kept under observation. It produced four clutches of eggs from10 May to 22 June. The average time interval of the successive clutches was 14.75 days, and the average clutch size was 2.75 eggs. The production of the successive clutches without mating in captivity may be due to the sperm storage in the female reproductive tract. The fertilization ratio and hatching ratio were 100% for the first three clutches, but only 33.3% and none, respectively, for the fourth clutch. The clutch order was significantly positively correlated with egg width and also with egg weight. T. stejnegeri was an intermediate egg retainer, whose gestation (egg retention) period was at an average of 25% of the total embryonic development period.

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Based on 20,000 records representing c. 11,000 individuals from an 8-year capture-mark-recapture (CMR) study, we tested and confirmed a new case of invariant clutch size (ICS) in a sexually dichromatic lacertid lizard, Takydromus viridipunctatus. In the grassland habitat of the early succession stage, females showed strictly low and invariant clutch size, multiple clutches in a breeding season, high reproductive potential, and annual breeding cycles that correspond to the emergence of male courtship coloration. The hatchlings mature quickly, and join the adult cohort for breeding within a few months, whereas adults show low survival rates and a short lifespan, such that most die within one year. Mortality increased in both sexes during the breeding season, especially in females, indicating an unequal cost of reproduction in survival. These life history characters may be explained by two nonexclusive hypotheses of ICS—arboreal hypothesis and predation hypothesis—within the ecological context of their habitat. Our study highlights a confirmed case of ICS, which adapts well to this r-selected grassland habitat that experiences seasonal fluctuation and frequent disturbance.

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Caudal autotomy in lizards has intrigued scientists for more than 100 years. Because of the relative lack of literature under natural conditions, the complicated association among field autotomy rate, real predation pressure, the long-term cost of tail loss, and the benefit of regeneration remains equivocal. In this study, we conducted a 7-year capture–mark–recapture (CMR) programme with a wild population of a sexually dichromatic lizard, Takydromus viridipunctatus. We used autotomy indexes and a contemporary bird census mega-dataset of four predatory birds as predictors to examine the association between tail loss and predation pressure. We further estimated the survival cost of tail loss and alleviation by regeneration under natural conditions through CMR modelling. We found that large and small avian predators affect lizard survival through the following two routes: the larger-sized cattle egret causes direct mortality while the smaller shrikes and kestrels are the major causes of autotomy. Following autotomy, the survival rate of tailless individuals over the next month was significantly lower than that of tailed individuals, especially males during the breeding season, which showed a decline of greater than 30%. This sex-related difference further demonstrated the importance of reproductive costs for males in this sexually dichromatic species. However, the risk of mortality returned to baseline after the tails were fully grown. This study indicates the benefit of tail regeneration under natural conditions, which increases our understanding of the cost–benefit dynamics of caudal autotomy and further explains the maintenance of this trait as an evolutionarily beneficial adaption to long-term predator–prey interactions.

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Notes on the corrected indentification and redescription of a ground lizard, Platyplacopus kuehnei, with a revised key of the family Lacertidae from Taiwan. Bull. In st. Zool. Academia Sinica 19(2): 63-65. The measurements and description of a new specimen of the Formosan ground lizard, Platyplacopus kuehnei (Van Denburgh) collected from Taipei in 1977, were given. Platyplacopus kuehnei which has been mistakenly named Takydromus kuehnei since 1909 should be rectified. A revised key is given for the family Lacertidae in Taiwan.

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Twelve tetra-repeated microsatellite loci were developed for the island-endemic grass lizard Takydromus formosanus (Squamata: Lacertidae). We characterized these loci by genotyping 51 individuals sampled from six localities. The number of alleles per locus ranged between 12 and 23, whereas the observed heterozygosity ranged between 0.686 and 0.941. None of these loci showed gametic disequilibrium or sex linkage. With such high polymorphism, we believe that these loci should be suitable for fine-scaled analysis of population structure, hybridizations or individual identification of grass lizards.

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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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Pliable-shelled eggs of Takydromus septentrionalis increased in wet mass over the incubation due to absorbing water from the substrate.The levels of substrate moisture significantly influenced final masses of the eggs incubated at 26 and 28℃.Eggs gained more mass in wetter substrates than in drier substrates.In contrast to those at 26 and 28℃,final masses of the eggs incubated at 24℃ were independent of the level of substrate moisture.Due to the positive relationship between final and initial egg masses seen in all cases,variations in final egg mass could be partly explained by the variation in initial egg mass.Within the range from 24 to 32℃,incubation temperature significantly influenced duration of incubation,hatching success,and size,mass,and sprint speed of hatchlings,with the eggs incubated at 24 and 26℃ exhibiting higher hatching successes and producing larger,heavier,and faster-running hatchlings.Within the range from -220 to 0 kPa,incubation hydric environments had no impacts on duration of incubation,hatching success,and size,mass,and sprint speed of hatchlings.Incubation thermal and hydric environments did not determine sex in T.septentrionalis.

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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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1. Warming temperatures caused by climate change are predicted to vary temporally and spatially. For mid- and high-latitude reptiles, the seasonal variation in warming temperatures experienced by embryos and hatchlings may determine offspring fitness, yet this has remained largely unexplored. 2. To evaluate the independent and interactive influence of seasonal variation in warming temperatures on embryonic and hatchling development, we incubated eggs and reared hatchlings of a cold-climate oviparous ectothermic species, the Heilongjiang grass lizard (Takydromus amurensis), following a 2×2 factorial design (present climate versus warming climate for embryos × present climate versus warming climate for hatchlings). We then evaluated embryonic and hatchling development, including hatching success, incubation period, initial hatchling body size, hatchling metabolic rate, growth rate and survival in the mesocosms. 3. We found that warming temperatures shortened the incubation period and produced hatchlings with higher survival rates than those incubated under the present climate conditions. Similarly, hatchlings reared under a warming climate had similar growth rates and resting metabolic rates, but higher survival rates than those reared under the present climate. Hatchlings that experienced both warming incubation and warming growth conditions had the highest survival rates. 4. This study revealed that moderate warming temperatures (Representative Concentration Pathway, RCP 4.5, 1.1–2.6°C) experienced by embryos and hatchlings interact to benefit hatchling fitness in cold-climate oviparous ectotherms. Our study also highlighted the importance of integrating seasonal variation in warming temperatures when evaluating the responses to climate warming in multiple developmental stages in oviparous ectotherms.

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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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年,在對山東省膠東半島最大的山脈--昆崳山進行兩棲爬行動物的普查中,采到2號活體蜥蜴標本,經鑒定為蜥蜴科草蜥屬北草蜥(Takydromus septentronalis Günther),是山東省爬行動物新記錄,現報道如下.

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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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Sexual size dimorphism (SSD) has long attracted the attention of biologists, and life-history variation is thought to play an important role in the evolution of SSD. Here we quantified SSD and female reproductive traits to identify potential associations between SSD and female reproduction in the white-striped grass lizard Takydromus wolteri. In a population from Chuzhou, China, the largest male and female were 53.0 mm and 57.5 mm in snout-vent length (SVL), respectively. Females were larger in SVL and abdomen length, whereas males were larger in head size and tail length. Females produced up to five clutches of eggs during the breeding season, with large females producing more clutches and more eggs per clutch than small ones. As a result, large females had a higher annual fecundity and reproductive output. Egg size was positively correlated with maternal SVL in the first clutch, but not in subsequent clutches. These results suggest that T. wolteri is a species with female-biased SSD, and that fecundity selection, in which large females have higher fecundity due to their higher capacity for laying eggs, is likely correlated with the evolution of SSD in this species

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We used the northern grass lizard Takydromus septentrionalis as a model animal to examine the energy allocation strategy and whether body temperature can affect maternal reproductive traits in lizards. We maintained adult females collected from the field soon after winter dormancy under three thermal conditions (24.0±1.0°C, 28.0±1.0°C and 32.0±1.0°C) throughout the subsequent reproductive season, and then recorded their energy intake, digestive efficiency, body size and mass changes, and reproductive parameters. Manipulation of body temperature in reproducing females affected energy intake and several maternal reproductive characteristics. Females allocated energy to reproduction following the fulfillment of energetic requirements for metabolism and growth. The reduced energy availability for reproduction constrained females at low or high body temperatures to produce fewer and smaller eggs per season but had no role in influencing the number of eggs produced in single reproductive bouts. Females used currently acquired energy to fuel reproduction, and this strategy did not shift seasonally from the first to subsequent clutches. Our study supports the idea that the use of currently acquired energy for reproduction may be more widespread in ectotherms than thought before, and shows for the first time that egg size is thermally plastic in T. septentrionalis, where eggs produced in single reproductive bouts are believed to be well optimized for size.

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We performed a common garden experiment to assess the differences in life history associated with local adaptation in Takydromus wolteri. Adults from Harbin (cold) and Chuzhou (warm) populations were maintained in common conditions, and their life-history and hatchling characteristics were compared. Harbin and Chuzhou females showed the same patterns of seasonal shifts in egg size and clutch size and produced eggs of similar size, but differed in all other reproductive characteristics. Harbin females laid larger and heavier clutches later and less frequently than Chuzhou females of the same body size. Harbin eggs hatched earlier and resulted in larger hatchlings with shorter tail, forelimb and hindlimb lengths. With respect to local adaptation of life history, we summarize our findings as follows. First, cold-climate females mature later, at a larger size, by prolonging growth and delaying reproduction because of thermal constraints on the time available for activity. Second, cold-climate females produce larger clutches in shorter time periods and invest more yolk in eggs to increase annual fecundity and reproductive output. Third, cold-climate eggs hatch earlier to compensate for a reduced growing season and earlier winter. Fourth, cold-climate hatchlings have a body shape associated with reduced heat exchange and maintenance costs, as predicted by Allen’s rule.

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In order to compare the geographic variation of the morphological and reproductive characteristics between different populations of Takydromus wolteri, this study was conducted between April and August, 2011 and 2012. Sexual size dimorphism (SSD) and female reproductive output of the Takydromus wolteri population from Harbin (Heilongjiang Province) were studied with the method of field capturing and laboratory feeding. Our results showed that the Harbin population of T. wolteri was in SSD and the female was significant larger than the male in snout-vent length (SVL) (F1,62=15.337, P<0.001). The mean SVL of females was 55.86 mm while the mean SVL of males was 52.13 mm. Females had larger abdomen length (F1,61=4.937, P< 0.05) and males had larger heads (head length: F1,61=103.56, P<0.001; head width: F1,61=68.524, P<0.001) as the same SVL because larger females had a higher annual fecundity (r=0.732,F1,18=20.821,P<0.01) and reproductive output (r=0.681,F1,18=15.579, P<0.01). SSD was positively geographic variation in the T. wolteri of different population, and SVL of Harbin population in the north was obviously larger than Chuzhou population in the south in both sexes, which accorded with the Bergman’s rule.

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The complete mitogenome sequence of Takydromus amurensis (Squamata: Lacertidae) is determined using long PCR for the first time in this study. It is a circular molecule of 17 333 bp in length (GenBank accession number: KU641018). Similar to the most other lizards, the complete mtDNA sequence of T. amurensis contained two rRNA genes (12S rRNA and 16S rRNA), 22 tRNA genes, 13 protein-coding genes (PCGs) and a control region (D-loop). The nucleotide composition was 31.23% A, 26.06% C, 13.91% G and 28.8% T. Mitochondrial genomes analyses based on NJ method yield phylogenetic trees, including 14 reported lizards belonging to three families (Lacertidae, Gekkonidae and Agamidae). These molecular data presented here provide a useful tool for systematic analyses of genus Takydromus.

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The ecology of most squamates from the Republic of Korea is poorly understood: information on tolerances to environmental variables, movement patterns, home range sizes, and other aspects of their natural history and ecological requirements are lacking. In turn, this lack of knowledge presents an obstacle to effective conservation management. Currently and at the national level, two of Korea’s eleven terrestrial snake species are listed as threatened or near threatened: Elaphe schrenckii and Sibynophis chinensis, and one out of the six lizard species (Eremias argus) is listed as threatened. However, various threats including habitat loss, climate change and poaching may have already but unknowingly elevated other Korean reptiles to threatened statuses. To help resource managers in developing conservation programs, we provide a summary of the literature on threats to Korean squamates, a national recommended threat status, and species accounts focused on Korean populations. We recommend listing Hebius vibakari, E. argus, Scincella huanrenensis and Gekko japonicus under a higher threat level than the one provided by either the Korean National Institute of Biological Resources or the IUCN Red List. Our results highlight that conservation plans are urgently needed for Korean squamates, mostly because of habitat destruction, and that additional research has to be conducted on most species as there is a clear need for integrated ecological studies and active monitoring programs.

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We present here details of a collection of herpetofauna from Bangladesh, held at the Jahangirnagar University Herpetological Group, Dhaka. This collection is comprised of at least 58 species, six are tentatively identi ed to their nearest described species which themselves are recognised to consist of more than one cryptic species. Four species could not be accurately identi ed to species level. It is likely that some of these ten “species” represent cryptic species and possibly undescribed taxa. Several of the localities presented here provide the rst con rmed localities for species previously listed as present in Bangladesh, but were not con rmed with locality or specimen data. Other species in the collection represent previously unknown localities, e.g., Euphlyctis hexadactylus, Uperodon globulosus and Cyrtodactylus sp. Eutropis multifasciata, and Hylarana cf. nigrovittata, are here recorded from Bangladesh for the rst time based on our collections and Takydromus khasiensis is also reported for the rst time from Bangladesh based on apparently unpublished specimens in the collection of the California Academy of Science.

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The work was conducted in Khankaiskii Nature Reserve in 2001—2006 years. It was found that Takydromus wolteri regularly occurs in the two parts of Nature Reserve: the fl oodplain of the Sungacha River (the part «Chorotovo boloto») and the floodplain of the Spassovka River (the part «Zhuravliny»). Data on new localities are presented.

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Data on the distribution of amphibians and reptiles in the Russian Far East in the reserves and national parks of federal importance are presented, as well as the information on the representation of these groups of animals in regional Red Data Books and the Red Data Book of the Russian Federation is provided.

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

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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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The collection of Amphibians and Reptiles of the Museum of Natural History of Ferrara has characteristics of historical representation, taxonomic and zoogeography that may be considered as examples of medium-sized types of collections more preva- lent in Italian museums. It consists of a historic part of the study and a donation from Guido Campadelli. The collection consists of 1739 specimens of amphibians (1734 of which held in 437 containers with ethanol and 5 taxidermized specimens) for a total of 78 amphibian species. There are 811 specimens of Reptiles (769 of which are stored in 526 containers with ethanol and 42 taxidermized specimens) representing a total of 108 species. The findings that form the collection of Amphibians come from 153 loca- lities of 15 states and 12 regions and 29 provinces of Italy. In the batrachological col- lection 62.1% of the total European species, and 94.3% of the total species in Italy, are represented. Zoogeographic regions most represented are the Palaearctic with 67% of the species of Amphibians in the collection and the Neotropical with 29%, fol- lowed by the Ethiopian (6%) and Nearctic (5%). Reptiles specimens come from 247 lo- calities of 15 states and 15 regions and 36 provinces of Italy. In the collection 46,2% of European species and 91,5% of the Italian ones are represented. The most represen- ted zoogeographic region are the Palaearctic with 89% of species in the collection, followed by the Ethiopian (14%), the Eastern (13%), Neotropical (11%) and the Nearctic (7%); australian and malagasy species do not exceed 3%. The herpetology collection was the subject of taxonomic and biogeographic analysis.

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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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Aim We provide a new quantitative analysis of lizard reproductive ecology. Comparative studies of lizard reproduction to date have usually considered life-history components separately. Instead, we examine the rate of production (productivity hereafter) calculated as the total mass of offspring produced in a year. We test whether productivity is influenced by proxies of adult mortality rates such as insularity and fossorial habits, by measures of temperature such as environmental and body temperatures, mode of reproduction and activity times, and by environmental productivity and diet. We further examine whether low productivity is linked to high extinction risk. Location World-wide. Methods We assembled a database containing 551 lizard species, their phylogenetic relationships and multiple life history and ecological variables from the literature. We use phylogenetically informed statistical models to estimate the factors related to lizard productivity. Results Some, but not all, predictions of metabolic and life-history theories are supported. When analysed separately, clutch size, relative clutch mass and brood frequency are poorly correlated with body mass, but their product – productivity – is well correlated with mass. The allometry of productivity scales similarly to metabolic rate, suggesting that a constant fraction of assimilated energy is allocated to production irrespective of body size. Island species were less productive than continental species.Mass-specific productivity was positively correlated with environmental temperature, but not with body temperature. Viviparous lizards were less productive than egg-laying species. Diet and primary productivity were not associated with productivity in any model. Other effects, including lower productivity of fossorial, nocturnal and active foraging species were confounded with phylogeny. Productivity was not lower in species at risk of extinction. Main conclusions Our analyses show the value of focusing on the rate of annual biomass production (productivity), and generally supported associations between productivity and environmental temperature, factors that affect mortality and the number of broods a lizard can produce in a year, but not with measures of body temperature, environmental productivity or diet.

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Aim: Small geographic ranges make species especially prone to extinction from an- thropogenic disturbances or natural stochastic events. We assemble and analyse a comprehensive dataset of all the world’s lizard species and identify the species with the smallest ranges—those known only from their type localities. We compare them to wide-ranging species to infer whether specific geographic regions or biological traits predispose species to have small ranges. Location: Global. Methods: We extensively surveyed museum collections, the primary literature and our own field records to identify all the species of lizards with a maximum linear geo- graphic extent of <10 km. We compared their biogeography, key biological traits and threat status to those of all other lizards. Results: One in seven lizards (927 of the 6,568 currently recognized species) are known only from their type localities. These include 213 species known only from a single specimen. Compared to more wide-ranging taxa, they mostly inhabit relatively inaccessible regions at lower, mostly tropical, latitudes. Surprisingly, we found that burrowing lifestyle is a relatively unimportant driver of small range size. Geckos are especially prone to having tiny ranges, and skinks dominate lists of such species not seen for over 50 years, as well as of species known only from their holotype. Two- thirds of these species have no IUCN assessments, and at least 20 are extinct. Main conclusions: Fourteen per cent of lizard diversity is restricted to a single location, often in inaccessible regions. These species are elusive, usually poorly known and little studied. Many face severe extinction risk, but current knowledge is inadequate to properly assess this for all of them. We recommend that such species become the focus of taxonomic, ecological and survey efforts.

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Thermal reaction norms of sprint speed were examined in three lacertid lizard species (Takydromus tachydromoides, T. smaragdinus, and T. dorsalis). We found inter- and intraspecific variations in maximum sprint speed, optimal body temperature, and thermal performance breadth for the best sprints. The thermal performance breadth of T. smaragdinus was broader than that of T. tachydromoides or T. dorsalis, whereas T. dorsalis sprinted faster than the others. In T. smaragdinus, individuals with narrower performance breadths run faster within their thermal range of expertise. There were “specialist” individuals of which performance depended heavily on high temperatures, as well as “generalist” individuals that performed well over a broad range of temperatures even within a single population. We discussed that the spatiotemporal instability of habitats might favor more than one type of thermal reaction norm of sprint performance, and this phenomenon may be driven by specialist–generalist trade-offs.

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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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Behavioral responses of the Japanese lacertid lizard,Takydromus tachydromoides, to its natural predator, the Japanese striped snake,Elaphe quadrivirgata, were examined under laboratory conditions.T. tachydromoides often thrashed its tail rapidly from side to side for short intervals (less than 0.5 s), usually just before fleeing away from an approaching snake. This tail vibration behavior appears to distract the snake away from the vulnerabl parts of the body to the tail, as tail vibration occurred more frequently when the head of the snake was closer to the lizard`s head than its tail. Distraction of snake attack by this action is suggested to be an antipredator tactic, because it was demonstrated that the vibration increased the chance of escape.

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Frequencies of movement, as an index of propensity for immobility, of Takydromus tachydromoides were measured in the encounters with a potential predator Elaphe quadrivirgata (Session A) or a sympatric non-predator Eumeces latiscutatus (Session B), or in the absence of other animals (Session C). Frequency of movement in Session A was significantly lower than in Sessions B and C. In Session B, T. tachydromoides moved significantly less frequently during the first 10min than during the latter 10min. The lizards previously exposed to E. quadrivirgata reduced their movements in the subsequent Session B with significantly high frequency when compared with animals without such experience. It is suggested that immobility of T. tachydromoides is an adaptive antipredator behavior to avoid detection by a visually orienting predator.

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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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Animals possess systems for sensing environmental temperature using temperature-sensitive ion channels called transient receptor potential channels (TRPs). Various TRPs have been identified and characterized in mammals. However, those of ectotherms, such as reptiles, are less well studied. Here, we identify the V subfamily of TRP (TRPV) in two reptile species: Japanese grass lizard (Takydromus tachydromoides) and Japanese four-lined ratsnake (Elaphe quadrivirgata). Phylogenetic analysis of TRPVs indicated that ectothermic reptilian TRPVs are more similar to those of endothermic chicken and mammals, than to other ectotherms, such as frog and fish. Expression analysis of TRPV4 mRNA in the lizard showed that its expression in tissues and organs is specifically controlled in cold environments and hibernation. The mRNA was ubiquitously expressed in seven tissues/organs examined. Both cold-treatment and hibernation lowered TRPV4 expression, but in a tissue/organ-specific manner. Cold-treatment reduced TRPV4 expression in tongue and muscle, while in hibernation it was reduced more widely in brain, tongue, heart, lung, and muscle. Interestingly, however, levels of TRPV4 mRNA in the skin remained unaffected after entering hibernation and cold-treatment, implying that TRPV4 in the skin may act as an environmental temperature sensor throughout the reptilian life cycle, including hibernation. This is the first report, to our knowledge, to describe reptilian TRPV4 in relation to hibernation.

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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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Based on a new herpetological collection from Ha Giang Province, Vietnam we report seven new records of amphibians and reptiles, comprising two species of Rhacophoridae (Rhacophorus kio, R. rhodopus), one species of Agamidae (Pseudocalotes brevipes), one species of Lacertidae (Takydromus kuehnei), one species of Colubridae (Lycodon futsingensis) and two species of Viperidae (Deinagkistrodon acutus, Protobothrops maolanensis). Our findings bring the species number of amphibians and reptiles recoded from Ha Giang Province to 110.

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Two exampls of double-tailed lizards, Takydromus tachydromoides SCHLEGEL and Anolis c. carolinensis, were examined. The Takydromus possessed a normally regenerated tail laterally, and an abnormally regenerated tail ventrally. X-ray photography showed the 38th vertebra bent ventrally at an angle of 90. Both the vertebra and the core of the ventral tail stained red with Mall`s Arizalin red-S, indicating that the core was formed of cartilage, or chondrin, or cartilagious tissue. This tail is presumed to have arisen from a broken bone. The regenerated tail of the Anolis was normal, without any Vertebrae.

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A morphological study on the monstered tail of two species of Lacertilia was done with X-ray photograph and Mall`s alizalin red-S staining method. One of them, Gekko japonicus, has a forked tail, and the other one, Takydromus tachydromoides, has a triple tail. The former`s regenerated tail develops at the level of the left transverse process of the 50th vertebra. It shows normal feature in structure. The latter`s primary regenerated tail develops at the caudal part of the 33rd vertebra and the secondary re- generated tail develops at the middle part of the primary regenerated tail. It seems to be constituted with cartilage or chondrin. These specimen were collected in Aichi Prefecture.

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A study of 3 spp. of the Lacertidae showed the chromosome number of Takydromus tachydromoides and T. smaragdinus as 2N = 38, each sp. having a pair of m-chromosomes of minute size; and of Lacerta vivipara as 2N = 36 with no m-chromosomes. A single specimen of T. smaragdinus possessed exceptionally large m-chromosomes, whereas they were of medium size in other specimens and extremely small in T. tachydromoides. Evidence of diminution in the size of the m-chromosomes by species and the assumption of their final disappearance may explain the numerical relation of the chromosomes found in Takydromus and L. vivipara.

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The Kühne’s Grass Lizard (Takydromus kuehnei) is a poorly studied indigenous species in Taiwan. In this report we incorporate additional information concerning reproduction with our previous observations to provide a more comprehensive description of reproduction in T. kuehnei from southwestern Taiwan. We collected 48 T. kuehnei (18 male, 19 female) from a secondary forest in Santzepu, Sheishan District, Chiayi County, as part of a herpetofauna survey in the area. We also obtained five clutches of eggs from these lizards. The smallest reproductively active female was 49 mm snout-vent length (SVL). Oviposition took place from April to July. Clutch sizes ranged from one to two eggs with an average of 1.8. A histological examination of five museum specimens confirmed that females may produce multiple clutches in the same year. We obtained 11 eggs that had an average length, width, and mass of 10.6 mm, 6.1 mm, and 1.8 g, respectively. The average relative clutch mass was 16.7%. Five eggs successfully hatched after an average incubation period of 32.6 days. The hatchlings had an average SVL of 23.8 mm, total length of 51.4 mm, and mass of 0.3 g.

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Thirteen (male = 2; female = 10; juvenile = 1) Takydromus formosanus were collected on an ad hoc basis from an Areca catechu plantation in Santzepu, Sheishan District, Chiayi County, as part of a herpetofauna research in the area. The smallest reproductively active female had a snout-vent length (SVL) of 44 mm, and the smallest male with an enlarged tail base had a SVL of 41 mm. A histological examination and oviposition indicated that the female reproductive cycle commences in April and lasts until at least June. No clutch size bigger than two eggs was recorded in this study. Takydromus formosanus is a poorly studied species, and many aspects of the reproductive biology of this species merits further empirical study.

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In 1967, the author conducted a herpetolog cal survey of the islands of Niijima, Shikinejima, and Kozushima, which belong to the Izu island chain of Japan. The harpetological fauna of thes islands is very poor, and no amphibians occur, but the author confirmed the occurrence of 4 species of reptiles belonging to 3 families, as listed below. Colubridae 1. Elaphe quadrivirgata (Boie, 1826) (Niijima, Kozushima). 2. Elaphe climacophora (Boie, 1826) (Niijima, Kozushima). Scincdae 3. Eumeces okadae Stejneger, 1907 (Niijima, Shikinejima, Kozushima). Lacertidae 4. Takydromus tachydromoides (Schlegel, 1838) (Kozushima).

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The fine structure and immunocytochemical localization of serotonin in the cells of the receptor line were studied in the parietal eye and pineal organ proper of the Japanese grass lizard, Takydromus tachydromoides. Typical photoreceptor cells (PC) were the predominant cell type in the receptor line of the parietal eye, the outer segments of which had regular stacks of numerous disks similar to those of cones. The pineal organ contained relatively few PCs, which showed less well-developed outer segments than those of the parietal eye. In contrast, rudimentary photoreceptor cells (SRPC) accounted for the majority of receptor cells in the pineal organ. These cells were structurally characterized by whorl-like lamellar outer segments and numerous dense-cored vesicles (80–280 nm in diameter). A small number of SRPC were also found in the parietal retina, which were similar to those in the pineal organ. In the parietal–pineal complex, numerous mitochondria located in the PC were larger and rounder than those in the SRPC. In the PC, basal processes prossessed only synaptic ribbons, whereas in the SRPC some of these processes contained synaptic ribbons and others contained dense-cored vesicles, rarely having both. Serotonin-immunoreactive cells were found not only in the pineal organ but also in the parietal eye, which closely resembled the cells of the receptor line in their size and shape. Furthermore, on immunoelectron microscopy for serotonin using the protein A-gold technique, gold particles indicating serotonin-immunoreactive sites were restricted in the core of dense-cored vesicles in the SRPC of the pineal organ. Regional differences in the distributions of the PC, SRPC and serotonin-immunoreactivity were found in the parietal-pineal complex.

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The Japanese grass lizard, Takydromus tachydromoides, is a species endemic to Japan. While several studies have investigated the reproductive ecology of T. tachydromoides, the embryogenesis of this species has not been reported in detail. In the current study, we observed the external morphological characteristics of 247 embryo specimens that developed under a constant temperature (28°C) throughout the duration from oviposition to hatching. We identified 17 consecutive developmental stages for the postovipositional developmental process of this species based on the staging criteria that have been widely used for the description of embryogenesis of lizards. The youngest embryos on the day of oviposition corresponded to stage 26, which is a relatively early period in pharyngula stages, and juveniles hatched at stage 42, approximately 30 days after oviposition. The entire developmental sequence of key morphological features was shared with other species of Lacertidae, except for the timing of the beginning of the first body pigmentation. This is the first description of the complete sequence of postovipositional developmental stages for the oviparous species of Lacertidae, providing valuable information for further evolutionary developmental studies.

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Squamate reptiles rely on the vomeronasal organ for chemoperception of their environment. This organ functionally associates with several structures, such as the nasolacrimal duct and choanal groove, which are morphologically diverse among species. In this study, we investigated the morphological characteristics and embryogenesis of the vomeronasal organ and its associated structures in the Japanese Grass Lizard (Takydromus tachydromoides). Separation of the complex of the nasolacrimal duct from the vomeronasal organ was observed in this species, a characteristic morphological feature among squamates. Additionally, we integrated the morphological characteristics of the vomeronasal organ and associated structures among squamates, focusing on topological relationships among the structures. The parsimony and likelihood methods for ancestral state reconstruction suggested the parallelism of several morphological characteristics during the evolutionary history of squamates: loss or reduction of the choanal groove, fusion between the nasolacrimal duct and choanal groove, and separation of the nasolacrimal duct from the vomeronasal organ.

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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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This paper analyzes the phylogeographical patterns of amphibian and reptile lineages distributed in the Ryukyu islands south of the Tokara Gap on the basis of relevant distributional data and phylogenetic hypotheses hitherto published. Results indicate the numerical dominance of highly relict lineages in the central Ryukyus, and the occurences of a few more or less relict lineages in the Miyako Group as well. On the other hand, most species and subspecies endemic to the Yaeyama Group or to the Yaeyama and Miyako Groups are considered to be direct consequences of vicariance with adjacent regions. Three major hypotheses regarding the Cenozoic paleogeography of the central and the southern Ryukyus are examined in the light of the present results and a modified hypothesis is proposed accordingly.

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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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We incubated eggs of Takydromus wolteri at four costant temperature to assess influence of incubation temperature on size, morphology, and locomotor performance of hatchlings. All viable eggs incubated in mass over the course of incubation due to absorption of water from surroundings, the mass gain was dependent on initial egg mass and incubation temperature. The duration of incubation at 24, 27, 30, and 33℃ averaged 41.9, 30.6, 25.8, and 22.6d, respectively. Incubation temperature significantly affected hatching success and body wet mass, carcass dry mass, and residual yolk dry mass of hatchlings, but it did not affect sex ratio, total dry mass, snout-vent length, and tail length of hatchlings. Compared to those from 33℃, hatchlings from 24℃ and 27℃ were greater in wet body mass and carcass dry mass, with a smaller quantity of yolk remaining unutilized at hatching. Incubation temperature affected some norphological traits of hatchlings, with hatchlings from 24℃ and 27℃ having larger size-free head length and width than did those from 33℃, and hatchlings from 24℃ generally had smaller size-free tympanum diameter than did those from higher temperatures.

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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 area traversed by an animal during food gathering, mating, and other routine activities is termed the home range (HR), and its size is believed to reflect behavioral and physiological requirements (e.g., access to mates and food). We evaluated both biological and methodological factors that may affect estimated HR sizes of lizards. We compiled a database of nearly 500 published sex 3 population data sets. These yielded usable data for 108 populations from 60 species, all with both sexes represented. Initial analyses indicated pervasive sex differences, so sexes were analyzed separately with both conventional and phylogenetically based statistical methods. First, we conducted conventional analyses of covariance, which treat each data point as statistically independent. Second, we repeated these analyses while comparing members of two major clades represented in our database, Iguania and Autarchoglossa. Third, we repeated the analyses after designating smaller clades, nested within major clades. Fourth, we performed a fully phylogenetic analysis via independent contrasts and a composite phylogeny assembled from published studies. All analyses showed highly significant effects of both biological (body size, diet, habitat type) and most methodological (calculation method and minimum number of sightings per individual, but not study duration) factors and covariates tested. The conventional analysis comparing Iguania and Autarchoglossa en toto indicated a significant difference, as has been suggested in the literature. The analysis with nested minor clades, however, revealed differences among these but not between the two major clades. The analysis with phylogenetically independent contrasts indicated no statistically significant differences either among minor clades or between major clades. We also used our database to compare the scaling of HR area to body mass, using recently published allometric equations for field metabolic rates of lizards. The similarity of allometric slopes suggests that, interspecifically, HR size scales directly with energetic requirements. Intraspecifically, however, we found that males in both Autarchoglossa and Iguania consistently had larger HRs than did females. This difference, presumably related to the need to maximize access to females, may impose a ‘‘cost of reproduction’’ on males.

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Based on a new herpetological collection from Ha Giang Province, Vietnam we report seven new records of amphibians and reptiles, comprising two species of Rhacophoridae (Rhacophorus kio, R. rhodopus), one species of Agamidae (Pseudocalotes brevipes), one species of Lacertidae (Takydromus kuehnei), one species of Colubridae (Lycodon futsingensis) and two species of Viperidae (Deinagkistrodon acutus, Protobothrops maolanensis). Our findings bring the species number of amphibians and reptiles recoded from Ha Giang Province to 110.

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The paper discusses biotopical and altitudinal distribution of Takydromus amurensis Peters, 1881 and Takydromus wolteri Fischer, 1885 inhabiting the north-eastern Asia. These species demonstrate in this area the changes of biotopical preference. T. amurensis prefers to occupy the forest zone less actively than in the southern part of range, 46 vs. 91%, respectively. T. wolteri avoids open spaces and partially enters the forest zone (7%). Both T. amurensis and T. wolteri actively inhabit the light forest zone (15 and 17%, respectively) and anthropogenic region (39 and 76%, respectively), the zone being optimal for their inhabitancy. In the southern part of natural habitat (Republic of Korea), the light forest zone plays an important role in their biotopical distribution (9% T. amurensis and 36% T. wolteri), though it is less preferred when compared to north-eastern part of the range. Two sites in the forest zone with weakly anthropogenic disruption (one in Russia and the other in Republic of Korea) were revealed, where species symbiotopy was observed. In both cases, T. wolteri was a predominant species. For the first time the altitudinal distribution data for lizards of genus Takydromus has been presented, in particular, for Russia. T. amurensis was found in the range of 11 – 633 m a.s.l. and T. wolteri — 14 – 321 m a.s.l. Maximal amplitude of altitudinal distribution of studied species has been indicated for the light forest zone. On the territory of Korea, T. wolteri has been found to have wider range of vertical distribution 4 – 424 m a.s.l.

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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 karyotypes of Takydromus amurensis and T. wolteri from Heilongjiang Province were examined using medulla cells and the colchicine-hypotonic air-dry technique. The results showed that the number of diploid chromosomes in the two species was 38 and were composed of 18 pairs of euchromosomes and a pair of sex chromosome, including 17 pairs of telocentric chromosomes and a pair of spot chromosomes. The sex chromosome was of the type ZW. According to karyotypic characteristics, they are all primitive species within Lacertiformes.

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Although lizards are important hosts for hard ticks (Ixodidae), very few studies have been conducted in South Korea. To determine whether or not hard ticks can infest lizards endemic to South Korea, we examined 77 lizards of four species (Eremias argus, Sincella vandenburghi, Takydromus amurensis, and Takydromus wolteri) that were collected at 22 different sites between April and October 2010. We confirmed that all four lizard species can be infested by Ixodes nipponensis larvae or nymphs. Of the 62 E. argus examined, we found an average of 12.5 larvae on two lizards and an average of one nymph on one lizard. We found seven nymphs on one S. vandenburghi. We found an average of two nymphs on two of the five T. amurensis and an average of one nymph on four of the nine T. wolteri. Ixodes nipponensis larvae and nymphs were found most frequently on the foreleg axillae (87.8%), followed by the forelegs (7.3%), the eyelids (2.4%), and the ears (2.4%) of the lizards. To the best of our knowledge, this is the first report of I. nipponensis infestations of lizards endemic to South Korea.

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Sexual dimorphism is widespread among lizards and may cause variation in morphology or color between sexes. These variations are determined by ecological, environmental, and genetic factors. In this study, we examined sexual dimorphism in Sikkim Grass Lizards (Takydromus sikkimensis) of Sikkim-Darjeeling Himalaya, India. We investigated whether differences exist in morphology and colors between sexes, and we also explored the breeding biology of this lizard in both captivity and under natural conditions. We observed female-biased sexual dimorphism in size and shape with females having greater snout vent length (SVL), forelimb length (FLL), hindlimb length (HLL), length of fourth finger (LFF), and snout to arm length (SAL) than males. Males had larger head length (HL) and head width (HW) compared to females. Our findings showed female-biased sexual dimorphism to fecundity selection, where larger females have higher fecundity than smaller females. Additionally, male-biased sexual dichromatism was observed during the breeding season. The primary habitat of Sikkim Grass Lizard in the study region is agro-ecosystems mostly below 1,000 m elevation. These ecosystems are undergoing rapid changes due to developmental projects, which could jeopardize the long-term survival of this species. Therefore, conservation efforts are needed and should focus on preserving a mosaic of agro-ecosystems, especially in the lower middle hills in Sikkim-Darjeeling region of the Eastern Himalaya.

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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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While the stability of sex chromosomes is widely accepted in viviparous mammals and birds, ectothermic vertebrates are still largely viewed as having frequent turnovers in sex determining systems. Frequent changes in sex determining systems in ectotherms could be problematic for field ecological studies as well as for breeding programs, as molecular sexing across a phylogenetically widespread spectrum of ectothermic vertebrates would not be possible. However, we recently documented that sex determining systems in three important reptile lineages (caenophidian snakes, iguanas, and lacertid lizards) are in fact highly conserved. 2.We applied a new molecular procedure to identify sex within each of these three lineages (encompassing altogether around 4,000 species, i.e. nearly 50% of the recent species of reptiles). This technique uses quantitative PCR (qPCR) to compare copy numbers of genes specific for their respective Z (in caenophidian snakes and lacertids) and X (in iguanas) chromosomes between male and female genomes. The DNA samples required can be collected relatively non-invasively. Unlike molecular sexing based on repetitive elements, this technique can be easily applied to previously unstudied species of these lineages, as the number of copies of protein coding genes linked to the differentiated sex chromosomes is evolutionary highly conserved in each. 3.We suggest that qPCR-based molecular sexing using the comparison of gene copy number is a practical choice for non-model species of caenophidian snakes, iguanas and lacertids. Morever, it should also soon be available for other reptile lineages with differentiated sex chromosomes.

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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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The squamates are composed of many taxa, among which there is morphological variation in the vomeronasal organ (VNO). To elucidate the evolution of chemoreception in squamate reptiles, morphological data from the VNO from a variety of squamate species is required. In this study, the morphology of the VNO of the grass lizard Takydromus tachydromoides was examined using light and electron microscopy. The VNO consists of a pair of dome-shaped structures, which communicate with the oral cavity. There are no associated glandular structures. Microvilli are present on the apical surfaces of receptor cells in its sensory epithelium, as well as on supporting cells, and there are centrioles and ciliary precursor bodies on the dendrites. In addition to ciliated cells and basal cells in the non-sensory epithelium, there is a novel type of non-ciliated cell in T. tachydromoides. They have constricted apical cytoplasm and microvilli instead of cilia, and are sparsely distributed in the epithelium. Based on these results, the variation in the morphology of the VNO in scincomorpha, a representative squamate taxon, is discussed.

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In this paper, the evolution and ecology of directional asymmetry (DA) during the developmental trajectory (DT) is compared with that of its product, morphological DA (MDA). DT and MDA are calculated for two bilateral morphological scale characters of lizards, the number of subdigital lamellae beneath the fourth toe in 10 agamid and 28 lacertid taxa, and the number of rows of ventral scales in 12 lacertid taxa. MDA, the subtraction between left and right sides (classical measure of DA), is functional in adult animals. Results confirm the hypodiesis that, in DT, the regression parameters a (constant) and b (regression slope) of counts on the right side with those on the left describe a developmental process. No phylogenetic or environmental effects were observed on a and b, but analyses considering both a and b together show non-random phyletic patterns. Independent analyses deduced the same ancestral DT in Agamidae and Lacertidae. In Lacertidae, distance between pairs of taxa in a + b (standardized values) correlates positively with die phylogenetic distance between taxa. Phyletic trends in MDA are indirect, and due to the link of MDA with a + b. The MDA of species is more dissimilar in sympatry than in allopatry. The phyletic trends suggest evolution of DT, while die association of MDA witii sympatry suggests that ecological pressures shape MDA in adult animals. Evolution of DT is independent from tiiat of its product, MDA–adaptive determinism defines the result of, but not the mechanistic process of, development. Deterministic environmental processes define MDA, and deterministic evolutionary processes define the interactive result of a and b, but not each separately. According to circumstances, different DTs produce similar or different MDA, and a particular DT can produce different MDAs.

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Heterozygosity-fitness correlations (HFCs) provide insights into the genetic bases of individual fitness variation in natural populations. However, despite decades of study, the biological significance of HFCs is still under debate. In this study, we investigated HFCs in a large population of the sexually dimorphic lizard Takydromus viridipunctatus (Lacertidae). Because of the high prevalence of parasitism from trombiculid mites in this lizard, we expect individual fitness (i.e., survival) to decrease with increasing parasite load. Furthermore, because morphological asymmetry is likely to influence individuals` mobility (i.e., limb asymmetry) and male biting ability during copulation (i.e., head asymmetry) in this species, we also hypothesize that individual fitness should decrease with increasing morphological asymmetry. Although we did not formally test the relationship between morphological asymmetry and fitness in this lizard, we demonstrated that survival decreased with increasing parasite load using a capture-mark-recapture data set. We used a separate sample of 140 lizards to test the correlations between individual heterozygosity (i.e., standardized mean d2 and HL based on 10 microsatellite loci) and the two fitness traits (i.e., parasite load and morphological asymmetry). We also evaluated and excluded the possibility that single-locus effects produced spurious HFCs. Our results suggest male-only, negative correlations between individual heterozygosity and parasite load and between individual heterozygosity and asymmetry, suggesting sex-specific, positive HFCs. Male T. viridipunctatus with higher heterozygosity tend to have lower parasite loads (i.e., higher survival) and lower asymmetry, providing a rare example of HFC in reptiles.

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The order Squamata is a species-rich group of reptiles including the groups Sauria, Serpentes and Amphisbaenia. Several members of these groups have been extensively used as model organisms for a great variety of studies of different fields including ecology, behaviour and medicine. Such studies require a well-established phylogenetic framework to trace the evolution and diversity of the studied traits in the squamate tree of life. However, the accuracy of phylogenetic inference in Squamata is unbalanced: while the position of the main squamate groups and high level relationships have been successfully and consistently estimated across phylogenetic studies, the phylogenetic relationships of some groups have provided inconsistent results over the years. The accurate estimation of relationships between these groups might have been hindered by methodological artefacts of the phylogenetic inference due to limited data and analytical tools. For the last couple of decades, and even presently, phylogenetic inference has relied heavily on mitochondrial DNA as the main molecular marker as a result of its advantages, such as the lack of recombination, easy amplification and high evolutionary rate. While its use is based on sound reasons, some drawbacks and limitations have been highlighted, such as the overall small size, and the fact that all genes are linked thus it represents a single locus. To overcome this, the addition of nuclear DNA complements the use of mitochondrial DNA in phylogenetic inference. Within the nuclear DNA, some markers may present so little variation that it is insufficient to recover phylogenetic relationships, yet slow-evolving nuclear genes have been the most widely used markers in phylogenetic studies. On the other hand, it has been shown that the use of highly informative fast evolving nuclear genes, in combination with mitochondrial DNA, can provide resolution at different parts and depths of the evolutionary history of the species. Phylogenetic analyses of multiple loci have historically been based on the concatenation approach, .i.e. the combination of the sequences into a single alignment that is then analysed to represent the species tree. This method, however, relies on the assumption that the most commonly occurring gene tree is equivalent to the true species tree of any taxa, which may not be true in all cases. Another limitation of the concatenation approach is the elevated percentage of missing data that it sometimes entails, especially in large taxon sets, meaning that the sequence representation is The order Squamata is a species-rich group of reptiles including the groups Sauria, Serpentes and Amphisbaenia. Several members of these groups have been extensively used as model organisms for a great variety of studies of different fields including ecology, behaviour and medicine. Such studies require a well-established phylogenetic framework to trace the evolution and diversity of the studied traits in the squamate tree of life. However, the accuracy of phylogenetic inference in Squamata is unbalanced: while the position of the main squamate groups and high level relationships have been successfully and consistently estimated across phylogenetic studies, the phylogenetic relationships of some groups have provided inconsistent results over the years. The accurate estimation of relationships between these groups might have been hindered by methodological artefacts of the phylogenetic inference due to limited data and analytical tools. For the last couple of decades, and even presently, phylogenetic inference has relied heavily on mitochondrial DNA as the main molecular marker as a result of its advantages, such as the lack of recombination, easy amplification and high evolutionary rate. While its use is based on sound reasons, some drawbacks and limitations have been highlighted, such as the overall small size, and the fact that all genes are linked thus it represents a single locus. To overcome this, the addition of nuclear DNA complements the use of mitochondrial DNA in phylogenetic inference. Within the nuclear DNA, some markers may present so little variation that it is insufficient to recover phylogenetic relationships, yet slow-evolving nuclear genes have been the most widely used markers in phylogenetic studies. On the other hand, it has been shown that the use of highly informative fast evolving nuclear genes, in combination with mitochondrial DNA, can provide resolution at different parts and depths of the evolutionary history of the species. Phylogenetic analyses of multiple loci have historically been based on the concatenation approach, .i.e. the combination of the sequences into a single alignment that is then analysed to represent the species tree. This method, however, relies on the assumption that the most commonly occurring gene tree is equivalent to the true species tree of any taxa, which may not be true in all cases. Another limitation of the concatenation approach is the elevated percentage of missing data that it sometimes entails, especially in large taxon sets, meaning that the sequence representation is Miocene and supported a scenario of east-to-west diversification. The little morphological and phylogenetic evidence for the distinctiveness between Rhinechis and Zamenis supports a classification lumping which better reflects their evolutionary history and, based on the priority rule, R. scalaris is moved into the genus Zamenis and designated as Zamenis scalaris comb. nov. Within the Gallotiinae subfamily, the combined use of fast nuclear markers and the species tree was fruitful for the inference of a robust phylogeny of the genus Psammodromus, distributed in the Iberian Peninsula and North Africa. In this study we analysed for the first time the six Psammodromus species to infer the association between the main cladogenetic events within this genus and the complex biogeographic dynamics across the Strait of Gibraltar. The inferred phylogeographical history suggests that Psammodromus probably originated in Iberia. The African species and the African lineage of P. algirus were a result of two over-sea dispersal events towards Africa that occurred 10 Ma and 1.5 Ma, while continental vicariance events might have shaped the diversification of the species within Iberia and Africa. These results, combined with previous literature, provide compelling evidence that major biotic exchanges occurred across the Strait of Gibraltar well before or long after the land connection during the Messinian Salinity Crisis (5.9 – 5.33 Ma). These findings suggest caution in the application of the relatively short event of the opening of the Strait of Gibraltar at the end of the Messinian Salinity Crisis as a cause for divergence in molecular clock calibrations, which is a common approach in literature. At the species level, the use of fast evolving nuclear markers proved to be fruitful to understand the geographical pattern of phyletic diversification within the Omanosaura lizards belonging to the Eremiadini tribe, endemic to the Hajar Mountains in the Arabian Peninsula. Multilocus phylogenetic analyses recovered two highly divergent lineages within O. cyanura which are geographically associated to the northernmost and to the south and eastern regions of the Hajar Mountains. These lineages are reciprocally monophyletic at both mitochondrial and nuclear loci suggesting a long history of independent evolution and the need of a comprehensive taxonomic assessment. Omanosaura cyanura represents an additional case of cryptic diversity in the north Hajar Mountains, an area that has been demonstrated to hide high levels of genetic diversity in other reptile groups. Overall, the work developed in this dissertation has demonstrated that the inclusion of fast evolving nuclear genes and the coalescent species tree approach was relevant for resolving challenging phylogenetic questions on selected squamate groups. This approach allowed the recovery of new clades and provided support for old and recent relationships. These methods have further allowed the comparison between contrasting phylogenetic hypotheses on the Lacertini and Colubrinae, demonstrating that the inference based on the supermatrix approach used by many recent studies may provide high support for incorrect nodes. Future research directions include (i) the application of the phylogenomic approach based on Next Generation Sequence data to further resolve basal polytomy within the Lacertini and Colubrinae radiations; (ii) a taxonomical assessment of Omanosaura; (iii) a re-evaluation of molecular clock calibrations based on the end of the Messinian Salinity Crisis as general cause for divergence, in order to account for biotic exchanges by over-sea dispersal across the Mediterranean well before or long after the land connection during this period, as suggested by emerging literature and the study case on Psammodromus.

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In order to characterize the current status and distribution of reptiles currently inhabiting the Republic of Korea, the results of survey data were analyzed in this study. The results of this analysis showed that the reptiles were distributed into two orders, seven families, and 21 species, with the exception of six marine reptiles, including Dermochelys coriacea schlegelii, Chelonia mydas japonica, Caretta caretta, Hydrophis melanocephalus, Hydrophis cyanocinctus, and Pelamis platrrus. Also, the taxonomic revisions of several species were suggested, although this would be ill-advised before further studies can be conducted regarding the current status of marine reptiles in the region.

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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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A survey of reptile-associated ticks and their infection status with severe fever with thrombocytopenia syndrome (SFTS) virus was conducted to determine the relative abundance and distribution among lizards, skinks, and snakes in the Republic of Korea (ROK). In total, 132 reptiles, including 49 lizards (two species), 15 skinks (one species), and 68 snakes (eight species) were collected. In total, 84 ixodid ticks belonging to two genera (Ixodes and Amblyomma) were collected from 28/132 (21.2%) lizards, skinks, and snakes. Ixodes nipponensis Kitaoka & Saito was only collected from lizards and skinks, while Amblyomma testudinarium Koch was only collected from snakes. Takydromus wolteri had the highest tick index (0.7; total number ticks/total number collected hosts) among lizards and skinks, while Rhabdophis tigrinus had the highest tick index (2.2) among the snakes. Ixodes nipponensis larvae and nymphs accounted for 11.1% and 88.9%, respectively, of all ticks collected from lizards and skinks, while only A. testudinarium nymphs were collected from snakes. Nymphs of both species of ticks were collected from lizards and skinks from April to October, while I. nipponensis larvae were collected only from September to October. Ixodes nipponensis larvae and nymphs were preferentially attached to the lateral trunk (83.3%) and the foreleg axillae (16.7%) of lizards and skinks. SFTS virus was detected in both I. nipponensis and A. testudinarium collected from lizards and snakes. Phylogenetic analysis of SFTS viruses of ticks collected from two lizards and one snake demonstrated close relationships with SFTS virus strains observed from humans and ticks in the ROK, China, and Japan. These results implicate lizards and snakes as potential hosts of SFTS virus.

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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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Life history traits may vary within and among species. Rarely, however, are both variations examined concurrently to identify the life history adaptation. We found that female body size, offspring number and size, and incubation period showed convergent evolution in two lacertid lizards (Takydromus wolteri and Eremias argus) that occur sympatrically in high-latitude and low-latitude localities. Females from the high-latitude population were larger and produced larger clutches than those from the low-latitude population. In both species, the incubation period was shorter for the high-latitude population than for the low-latitude population. However, the physiological mechanism underlying the shorter incubation period differed between the species. These results suggest that: (1) sympatric lizards may adopt similar reproductive strategies in response to their common environments, and (2) embryonic development of the two species follows different pathways for adaptation to low temperatures. This study highlights the importance of understanding the adaptive evolution of life history in response to environmental changes at the embryonic life stages.

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Parental effects may produce adaptive or maladaptive plasticity that either facilitates persistence or increases the extinction risk of species and populations in a changing climate. However, empirical evidence of transgenerational adaptive plastic responses to climate change is still scarce. Here we conducted thermal manipulation experiments with a factorial design in a Chinese lacertid lizard ( Takydromus septentrionalis) to identify the fitness consequences of parental effects in response to climate warming. Compared to present climate conditions, a simulated warming climate significantly advanced the timing of oviposition, depressed the immune capability of post-partum females, and decreased the hatching success of embryos, but did not affect female reproductive output (clutch size and egg mass). These results indicate that maternal warming negatively affects female health, and embryonic hatchability. More interestingly, we found that offspring from parents exposed to warming environments survived well under a simulated warming climate, but not under a present climate scenario. Accordingly, our study demonstrates anticipatory parental effects in response to a warming climate in an ectothermic vertebrate. However, the fitness consequences of this parental effect will depend on future climate change scenarios.

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A new grass lizard, Takydromus toyamai, is described from the Miyako Island Group, Ryukyu Archipelago. The Miyako population has been considered to be conspecific with T. smaragdinus from the Okinawa, Amami, and Tokara Groups of the archipelago, but it is differentiated from populations of these three island groups by having a greater number of ventral rows (eight) and distinctly keeled temporal scales, and in lacking a white ventrolateral line on head and body. The definition of T. smaragdinus is revised accordingly and a lectotype is designated. A dichotomous key is provided for the Japanese and Taiwanese species of the genus Takydromus.

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We investigated the genetic population structure of the Japanese grass lizard, Takydromus tachydromoides, based on polymerase chain reaction—restriction fragment length polymorphisms (PCR-RFLPs) and DNA sequence analyses of the mitochondrial cytochrome b gene. The PCR-RFLP analysis of 163 specimens collected from the main islands of Japan and the adjacent islands identified a total of 25 mitochondrial haplotypes (mitotypes). The phylogeny of the mitotypes revealed that Osumi and Tokara Island populations were remarkably diverged from all other populations on the main islands of Japan. Furthermore, several regional groups were also recognized at the western part of Japan, namely mitotype groups in Kyushu, those in the western Honshu, and those in the central Honshu and Shikoku. In contrast, little genetic diversity was observed throughout eastern Japan. The regional genetic differentiation and recent range expansion of this species are considered to be associated with past geological events and climate changes.

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Growth of Takydromus tachydromoides was studied at Fukuoka in southwestern Japan. From the data of growth in SVL, the lizards were classified into three age classes. The growth rate of SVL did not differ between sexes. The mean body weight of adult lizards differed significantly between sexes, but SVL did not.

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The home range of Takydromus tachydromoides was studied in a grassland area from April 1977 to November 1978. The mean size of home range did not differ markedly between sexes; 136.5 m2 for males and 130.8 m2 for females. Home ranges of adults overlapped greatly in each sex, and the lizard was considered to be non-territorial. Individuals showed return movement to a definite area (sleeping site) within the home range, and the home range did not shift within a year or between years. Characteristics of the home range of this grassland-inhabiting lizard were discussed in relation to resource abundance and predation pressure.

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A field study of the Japanese grass lizard Takydromus tachydromoides was carried out at Mito in central Japan. The mean growth rate of yearling females was highest during late spring. They grew little during the reproductive season (mid-summer), but resumed growth somewhat in late summer. Most energy is apparently directed to body growth in yearling females before the first reproduction. Thereafter most energy is allocated to reproduction.

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Reproduction in females of the kanahebi lizard, Takydromus tachydromoides, was studied in Mito, Ibaraki Prefecture. Yearling females larger than 40mm SVL and not reproductive early in the reproductive season (April-May) might be judged to be mature because their largest unyolked follicles were the same size as those of reproductive females (1.3mm or larger). Most yearlings bred once or twice late in the reproductive season (June-July), and they had a mean clutch size of 3.0. Older females reproduced from April through July, and they had a mean clutch size of 4.2 for the first clutch and of 4.0 for the second and third. In this population the mean dry weight of oviducal eggs was 84mg, and the mean ratio of clutch weight to body weight was 0.729.

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The Chung-an ground lizard, Takydromus sylvaticus, long known only from five females collected in Fujian Province of China, was recently found in Anhui Province of the country. The newly collected specimens included the first male specimens (n=7), in which the tail bases were stouter than in conspecific females. Ventral scales in these males were invariably keeled, whereas in females those forming the four median rows were smooth. Females had more dorsal scales (47–48) than males (42–43) in the median longitudinal row between axilla and groin. Takydromus sylvaticus differed from the closely related T. dorsalis in having more dorsal scales in the median longitudinal row between axilla and groin (42–48 versus 32–40, respectively), more rows of large dorsal scales between the hind legs (10 versus 7–8, respectively), shorter hind legs, two distinct white longitudinal stripes on each side of the body, and no black stripe in the preocular and postocular regions. The number of transverse rows of ventral scales between collar and femoral pores in T. sylvaticus(26–28) was larger, but with a slight overlap, than that in T. dorsalis (22–26). Ecological observations on T. sylvaticus were also documented.

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The Chung-an ground lizard Takydromus sylvaticus is an endemic and long-lost reptile species in China. In this study, its mitochondrial genome (mitogenome) sequence was firstly determined by long PCR and primer walking methods. The genome is 17,838 bp long and contains 13 protein-coding genes, 22 transfer RNA genes, 2 ribosomal RNA genes and 2 main noncoding regions (the control region and the putative L-strand replication origin). The gene order and contents in the T. sylvaticus mitogenome is identical to that found in typical vertebrates, suggesting that it represents an ancestral arrangement. Within the control region, typical conserved domains and distinct repeat regions were identified.

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Climate change poses different threats to animals across latitudes. Tropical species have been proposed to be more vulnerable to climate change. However, the responses of animals from tropical mountains to thermal variation and climate change have been scarcely studied. Here, we investigated the thermal biology traits of a tropical lizard (Takydromus kuehnei) distributed at high elevations (>950 m) and evaluated the vulnerabilities of T. kuehnei by thermal biology traits, thermal safety margin, and thermoregulatory effectiveness. The average active body temperatures of T. kuehnei in the field were 26.28_C and 30.65_C in April and June, respectively. The selected body temperature was 33.23_C, and the optimal temperature for locomotion was 30.60_C. The critical thermal minimum and critical thermal maximum temperatures were 4.79_C and 43.37_C, respectively. Accordingly, the thermal safety margin (1.23_C) and thermoregulatory effectiveness (1.23_C) predicted that T. kuehnei distributed in tropical mountains were not significantly depressed by environmental temperatures. This study implies that high-elevation species in tropical regions may not be severely threatened by ongoing climate change and highlights the importance of thermal biology traits in evaluating the vulnerability of species 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 three year study of a population of Takydromus tachydromoides in central Honshu, Japan revealed that yolk deposition in follicles of juvenile lizards begins in the late spring following their first hibernation, at a body length exceeding 41 mm. Adult females emerge from hibernation with yolked follicles. Oviposition of first year lizards begins in early June, reaches a maximum in July and ends before mid-August. Older females begin oviposition in early May, reach their peak in the first half of June, and cease laying in late July. In experimental lizards corpora lutea regressed from an average of 2.1 mm on the day of oviposition to 1.0 mm five days postoviposition. These persist as corpora albicantia apparently for the life of the lizard. Estimates of age groupings by counts of corpora albicantia suggests that first year females contain none in spring and 4-13 in fall; second year lizards, 1-12 in spring and 14-26 in fall; third year females, 14-23 in spring and 26-39 in fall; and fourth year females, 31-40 in spring and 51-69 in fall. The incidence of corpora albicantia in adult lizards collected from 15 August until hibernation is 97%. Clutch size of first year lizards ranges from 1 to 4 for all clutches, with means of 2.6 for the first clutch and 2.4 for the second and third. Clutches of second year females range from 1 to 6, with means of 4.0 for the first and second, and 3.4 for the third clutch. Third year females lay clutches of 1-7 ova, with means of 4.4 and 5.1 for first and second clutches, respectively; third clutches approximate 3.5 eggs. Fourth year females apparently deposit 4-6 ova or more per clutch. Mean clutch number calculated by dividing average clutch size into the mean number of corpora albicantia in fall lizards is 2.9. A mean clutch number of 2.7 was obtained by dividing the length of the reproductive season by the time required to produce the first clutch. The reproductive season varied from 76 to 86 days for first year females, and from 112 to 127 days for older females. Reproductive potentials estimated by counts of yolked follicles, corpora lutea and oviductal eggs were 7.4, 11.4, and 12.8, for first, second and third year females, respectively. Estimated from the mean difference between counts of corpora albicantia in spring and fall lizards, they were 9.3, 12.9 and 11.6, in respective order. The latter method suggests a mean reproductive potential for fourth year females of 20.4. There is no difference in average clutch size between small and large females of the same age group, but second year females produce first clutches nearly twice as large as those of first year lizards. Average weights of individual eggs and total clutches increase as females age. Total clutch weight ranges from 30 to 50% of the females postovipositional weight. The sex ratio in all age groups in each season is 1:1. Of the total female sample, 62.5% had survived their first breeding season, 25.0% their second, 9.8% their third, and 2.7% their fourth season.

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easonal fluctuations in liver and fat body weights were studied in a population of Takydromus tachydromoides (Lacertidae) during two annual cycles. Liver and fat body weights are at minimum levels in late summer and rise rapidly in September and early October prior to hibernation. Livers of adult females and juveniles of both sexes show no apparent weight loss, but livers of adult males decrease in weight during hibernation, and continue to decline throughout spring and early summer. Livers of juvenile males and all females increase in weight during spring; those of adult females decrease during formation of clutches two and three in June and early July. Fat bodies of each age group of both sexes decrease in weight by approximately 50% during hibernation. Fat bodies of males continue to decline until July, although those of juvenile males increase immediately following hibernation. Adult and juvenile females increase fat reserves until vitellinogenesis for the first clutch increases sharply in the last half of April by adults and in late May by juveniles. Later clutches then exhaust remaining fat reserves. Fat bodies of adult females decreased in weight by 83 and 74% during formation of clutch one in 1966 and 1967. Age differences exist in proportionate organ weights of each sex during much of the season. During formation of clutch one, second year females had available approximately three times as much fat as did first year females.

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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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Vertebral laminae are bony ridges or sheets that connect important morphological landmarks on the vertebrae, like diapophyses or zygapophyses. They usually exhibit some serial variation throughout the column. A consistent terminology facilitates the morphological description of this variation, and the recognition of patterns that could be taxonomically significant and could serve as phylogenetic characters. Such a terminology was designed for saurischian dinosaurs, and has also been applied to other members of Archosauri- formes. Herein, this terminology is applied for the first time to lizards (Squamata). Probably due to their generally smaller size compared to saurischian dinosaurs, lizards have less developed vertebral laminae. Some laminae could not be recognized in this group and oth- ers require new names to account for differences in basic vertebral morphology. For instance, the fusion of diapophysis and parapophysis in lacertids into a structure called synapophysis necessitates the creation of the new term synapophyseal laminae for both diapophyseal and parapophyseal laminae. An assessment of occurrence and serial varia- tion in a number of lacertid species shows that some laminae develop throughout ontogeny or only occur in large-sized species, whereas the distribution of other laminae might prove to be taxonomically significant in future.

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Dating the time of divergence and understanding speciation processes are central to the study of the evolutionary history of organisms but are notoriously difficult. The difficulty is largely rooted in variations in the ancestral population size or in the genealogy variation across loci. To depict the speciation processes and divergence histories of three monophyletic Takydromus species endemic to Taiwan, we sequenced 20 nuclear loci and combined with one mitochondrial locus published in GenBank. They were analyzed by a multispecies coalescent approach within a Bayesian framework. Divergence dating based on the gene tree approach showed high variation among loci, and the divergence was estimated at an earlier date than when derived by the species tree approach. To test whether variations in the ancestral population size accounted for the majority of this variation, we conducted computer inferences using isolation-with-migration (IM) and approximate Bayesian computation (ABC) frameworks. The results revealed that gene flow during the early stage of speciation was strongly favored over the isolation model, and the initiation of the speciation process was far earlier than the dates estimated by gene- and species-based divergence dating. Due to their limited dispersal ability, it is suggested that geographic isolation may have played a major role in the divergence of these Takydromus species. Nevertheless, the current study reveals a more complex situation and demonstrates that gene flow during the speciation process cannot be overlooked and may have a great impact on divergence dating. By using multilocus data and incorporating Bayesian coalescence approaches, we provide a more biologically realistic framework for delineating the divergence history of Takydromus.

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Expression of nuptial color is usually energetically costly, and is therefore regarded as an ‘honest signal’ to reflect mate quality. In order to choose a mate with high quality, both sexes may benefit from the ability to precisely evaluate their mates through optimizing visual systems which is in turn partially regulated by opsin gene modification. However, how terrestrial vertebrates regulate their color vision sensitivity is poorly studied. The green-spotted grass lizard Takydromus viridipunctatus is a sexually dimorphic lizard in which males exhibit prominent green lateral colors in the breeding season. In order to clarify relationships among male coloration, female preference, and chromatic visual sensitivity, we conducted testosterone manipulation with mate choice experiments, and evaluated the change of opsin gene expression from different testosterone treatments and different seasons. The results indicated that males with testosterone supplementation showed a significant increase in nuptial color coverage, and were preferred by females in mate choice experiments. By using quantitative PCR (qPCR), we also found that higher levels of testosterone may lead to an increase in rhodopsin-like 2 (rh2) and a decrease in long-wavelength sensitive (lws) gene expression in males, a pattern which was also observed in wild males undergoing maturation as they approached the breeding season. In contrast, females showed the opposite pattern, with increased lws and decreased rh2 expression in the breeding season. We suggest this alteration may facilitate the ability of male lizards to more effectively evaluate color cues, and also may provide females with the ability to more effectively evaluate the brightness of potential mates. Our findings suggest that both sexes of this chromatically dimorphic lizard regulate their opsin expression seasonally, which might play an important role in the evolution of nuptial coloration.

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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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Shape variation in the vestibular system is often linked to microhabitat structure and locomotor performance. Highly circular and orthogonal semicircular canal pairs are linked to higher motion sensitivity. Here, we use 3D geometric morphometrics to investigate shape variation in the vestibular system within lacertid lizards and its relationship to balance control. We found that lacertids living in complex microhabitats possess narrow but longer vestibular systems, an S-shaped anterior canal, a straightened lateral canal and a short common crus. However, lacertids specialized for simple microhabitats (open areas) possess wider but shorter vestibular systems, more circular anterior and lateral canals, and a longer common crus. Contrary to our expectations, species living in simple microhabitats possess more anatomical adaptations that enhance the sensitivity of their vestibular system. This suggests that species inhabiting open areas may benefit from increased sensitivity given that they are potentially more visibile to predators and have lower shelter availability. Finally, the wider shape of the vestibular system of the open area species may be linked to a wider and potentially flattened skull, which may be related to sand-diving or prey hardness.

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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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A new species, Takydromus yunkaiensis J. Wang, Lyu, & Y.Y. Wang, sp. nov. is described based on a series of specimens collected from the Yunkaishan Nature Reserve located in the southern Yunkai Mountains, western Guangdong Province, China. The new species is a sister taxon to T. intermedius with a genetic divergence of 8.0–8.5% in the mitochondrial cytochrome b gene, and differs from all known congeners by a combination of the following morphological characters: (1) body size moderate, SVL 37.8–56.0 mm in males, 42.6–60.8 mm in females; (2) dorsal ground color brown; ventral surface green to yellow-green, but light blue-green on chin and throat, posteriorly green in adult males; (3) dorsolateral lines paired, strikingly yellowish-white bordered by black above and below, invisible or indistinct in juveniles and adult females; (4) flanks of body blackish brown with light brown marks in adult males; (5) presence of four pairs of chin-shields; (6) four supraoculars on each side; (7) presence of a row of supracilary granules that separate supracilaries from supraoculars; (8) two postnasals; (9) enlarged dorsal scales in six longitudinal rows on trunk of body, with strong keel; (10) enlarged ventral scales in six longitudinal rows, strongly keeled in males, smooth but outermost rows weakly keeled in females; (11) enlarged and keeled lateral scales in a row above ventrals; (12) femoral pores 2–3 on each side; (13) subdigital lamellae 20–23 under the fourth finger, 23–30 under the fourth toe; and (14) the first 2–3 subdigital lamellae under the fourth toe divided. The discovery of Takydromus yunkaiensis sp. nov. brings the total number of species of this genus to 24, of which nine occur in mainland China.

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We updated the checklists of extant, native amphibians and reptiles of China based on the previously published checklist of reptiles in 2015, the online checklist of amphibians on the database AmphibiaChina, newly published data as of December 2019, and previously uncollected literature prior to 2015. In total, the amphibian fauna of China consists of 515 species in 62 genera, 13 families, and three orders (Anura: 431 species in 47 genera and nine families; Caudata: 82 species in 14 genera and four families; Gymnophiona: one species in one genus and one family), while the reptilian fauna of China consists of 511 species in 135 genera, 35 families, and three orders (Crocodylia: one species in one genus and one family; Testudines: 34 species in 18 genera and six families; Squamata 466 species in 116 genera and 28 families [Serpentes: 256 species in 73 genera, 18 families; Lacertilia: 211 species in 43 genera and 10 families]). Specifically, for amphibians between 2015 and 2019, one family was recorded from China for the first time, two new genera were described, a genus was resurrected, a genus was recorded from China for the first time, 74 new, valid species were either described or resurrected, 18 recognized species were recorded from China for the first time, and six genera and eight species were considered as junior synonyms. For reptiles between 2015 and 2019, five subfamilies were elevated to the full family status, one new subfamily and a new genus were described, three genera were resurrected, three recognized genera were recorded from China for the first time, 35 new species were described, two species were resurrected from synonyms, six subspecies were elevated to the full species status, 10 recognized species were recorded from China for the first time, four genera and four species were considered as junior synonyms, and distribution records of one genus and four recognized species were removed from China. Furthermore, by reviewing literature before 2015, we make additional changes on the previous reptile checklist, including adding new records of three genera, elevating three subspecies to full species status, adding new records of three recognized species, synonymizing three genera and two species as junior synonyms, and removing the distribution record of a single recognized species from China. Lastly, we revise the Chinese common names of some reptilian groups with recomandations to maintain the stability of the Chinese common names. The number of new species and new national records for amphibians and reptiles between 2015 and 2019 in China accounts for 17.1% and 10.2% of the total number of species in each group, respectively. Because new species are described at considerable speed and given the constant changes in the taxonomy of China’s herpetofuna, it is crucial to update the checklists regularly and discuss the existing taxonomic problems, so that such information reflects the most current state of knowledge and are available for taxonomic researchers and conservation biologists alike.

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Following habitat fragmentation, the remnant faunal community will undergo a period of species loss or ‘relaxation.’ Theory predicts that species with particular life-history traits, such as a small population size, small geographical range, low fecundity and large body size, should be more vulnerable to fragmentation. In this study, we investigated the relationships between the above life-history traits and the fragmentation vulnerability index (the number of islands occupied) of five lizard species inhabiting recently isolated land-bridge islands in the Thousand Island Lake, China. Data on life-history traits were collected from field surveys (population density) and from the literature (body size, clutch size and geographical range size). The species–area relationships for lizards sampled from the mainland versus on the islands differed significantly (i.e. the number of species inhabiting islands was decreased relative to similar-sized areas on the mainland), indicating that species extinction has occurred on all of the study islands following isolation. For the fragmentation vulnerability index, model selection based on Akaike’s information criterion identified natural density at mainland sites as the best correlate of vulnerability to fragmentation, supporting the hypothesis that rare species are most vulnerable to local extinction and will be lost first from fragmented landscapes. In contrast, there was little evidence for an effect of lizards’ snout–vent length, clutch size or geographical range size on fragmentation vulnerability. Identification of species traits that render some species more vulnerable to fragmentation than others has important implications for conservation and can be used to aid direct management efforts.

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Many early descriptions of species of the genus Takydromus were based on limited diagnostic characteristics. This has caused considerable challenges in accurate species identification, meaning that a number of cryptic species have been erroneously identified as known species, resulting in substantially underestimated species diversity. We have integrated evidence from morphology and DNA sequence data to describe a new species of the Asian Grass Lizard, Takydromus albomaculosus sp. nov., based on two specimens from Tianjingshan Forestry Station, Ruyuan County, Guangdong Province, China. The new species can be distinguished from other known Takydromus species by distinctive morphological differences and significant genetic divergence in the mitochondrial COI gene. Morphologically, the new species differs from the known 22 congeners in the following combination of characters: relatively large body size, SVL 70.9 mm in female, 65.5 mm in male; presence of three or four pairs of femoral pores and four pairs of chin-shields; supraoculars three, rarely four, the posterior loreal either in contact with the anteriormost supraocular or not; the presence of supraciliary granules; the presence of six rows of ventral scales, strongly keeled in the male, and smooth but with the outermost row keeled in the female; the presence of enlarged and keeled lateral scales in a row above the ventrals, but shorter than the ventrals; subdigital lamellae 23–24 under the fourth finger and 29–30 under the fourth toe; dorsal surface brown, ventral surface green, flank dark brown or brown black with several prominent white round spots; the presence of a pair of white longitudinal dorsolateral stripes and a pair of white longitudinal ventrolateral stripes. Takydromus albomaculosus sp. nov. represents the twenty-third species of this genus, of which six species occur in northern Guangdong, where the biodiversity level of Takydromus is the second highest in the world, just below that of Taiwan.

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Abstract Background: Studies of lizards have shown that offspring size cannot be altered by manipulating clutch size in species with a high clutch frequency. This raises a question of whether clutch frequency has a key role in influencing the offspring size- number trade-off in lizards. Methodology/Principal Findings: To test the hypothesis that females reproducing more frequently are less likely to tradeoff offspring size against offspring number, we applied the follicle ablation technique to female Eremias argus (Lacertidae) from Handan (HD) and Gonghe (GH), the two populations that differ in clutch frequency. Follicle ablation resulted in enlargement of egg size in GH females, but not in HD females. GH females switched from producing a larger number of smaller eggs in the first clutch to a smaller number of larger eggs in the second clutch; HD females showed a similar pattern of seasonal shifts in egg size, but kept clutch size constant between the first two clutches. Thus, the egg size- number trade-off was evident in GH females, but not in HD females. Conclusions/Significance: As HD females (mean = 3.1 clutches per year) reproduce more frequently than do GH females (mean = 1.6 clutches per year), our data therefore validate the hypothesis tested. Our data also provide an inference that maximization of maternal fitness could be achieved in females by diverting a large enough, rather than a higher-than-usual, fraction of the available energy to individual offspring in a given reproductive episode.

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Three mantid species, Tenodera aridifolia (Stoll), Hierodura patellifera (Serville), and Statilia maculata Thunberg, of varying body size and habitats, exhibited defensive reactions when faced with a natural enemy, the Japanese skink, Takydromus tachydromoides (Schlegel). Defensive reactions were analyzed as primary or secondary defenses for all stages. Of the primary defenses, the cryptic reaction was the main defensive reaction: mantids lowered the prothorax and simultaneously stretched the prothoracic legs, the antennae were arranged forward, and the abdomen was sometimes extended posteriorly. The frequency of the subcryptic reaction as a primary defense tended to increase with more advanced developmental stage. The primary defensive strategy switched at specific developmental stage and also depended on the distance between the prey mantid and the natural enemy. In some instars, T. aridifolia and H. patellifera displayed the cryptic reaction when the mantid—foe distance was long, whereas they displayed the subcryptic reaction when the mantid—foe distance was short. However, S. maculata did not switch defense strategies based on the mantid—foe distance. The secondary defenses of young instar nymphs of T. aridifolia and H. patellifera were primarily defensive reactions, whereas the frequency of aggressive reactions tended to increase with developmental stage. S. maculata did not display aggressive reactions during any stage. We concluded that the primary and secondary defenses used by mantids depend on the developmental stage and mantid—foe distance. Moreover, three mantid species that vary in both body size and habitat showed different defensive reactions when faced with a natural enemy.