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

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The east Asian lacertid lizard genus,Takydromus, is well-defined but its relationships to other primitive Palaearctic lacertids are still uncertain and, although it has similarities to the lower Miocene Miolacerta, Takydromus does not appear to be identical with this. The 16 or 17 recognizable species are reviewed and a key provided for their identification; the remaining area of uncertainty about species boundaries is the Takydromus sexlineatus group in Assam. A phylogeny 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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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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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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From the perspectives of biodiversity conservation and management, there is an urgent need to have at hand current synopses of classification and distributions of species. In this paper, we review and summarize the classifications, Chinese and English names, type specimens, type localities and distributions of China’s lizard fauna to promote scientific exchange and species conservation among relevant people in China and internationally. As of December 31, 2020, a total of 230 species of lizard, belonging to 44 genera and 12 families, have been recorded from China, including 4 invasive species: 2 in the family Gekkonidae, 1 in Iguanidae and one in Dactyloidae. There are 115 endemic species, accounting for 50% of this group. The proportion of endemic species to China was the highest in the family Eublepharidae (84.62%), and the number of endemic species to China was the highest in the family Agamidae (42 species). The species distributions and classification in the “Fauna Sinica (Reptilia 2): Squamata (Lacertilia)” were updated. Among the provincial administrative units, there is a larger number of species in the southern provinces, such as Yunnan, Guangdong, Guangxi and Taiwan. Sichuan, Yunnan, and Taiwan harbour relatively more endemic species than others. A total of 154 species of lizards were first discovered in China, and the type localities of these species are concentrated in Yunnan, Taiwan, Tibet, Xinjiang, and Sichuan.

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

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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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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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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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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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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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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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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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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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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.