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

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

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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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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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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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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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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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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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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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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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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 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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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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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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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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We used Takydromus septentrionalis, a sexually size-monomorphic lacertid lizard, as a model system to test the hypothesis that sexual size monomorphism may evolve in lizards where reproductive performance is maximized at a similar body size for both sexes. We allowed lizards housed in laboratory enclosures to lay as many clutches (for females) as they could or to mate as many times (for males) as they could in a breeding season. Size-assortative mating was weak but evident in T. septentrionalis, as revealed by the fact that male and female snout–vent lengths (SVLs) in mating pairs were significantly and positively correlated. Mating frequency (indicative of male reproductive performance) varied from 1 to 8 per breeding season, generally increasing as SVL increased in adult males smaller than 67.4 mm SVL. Clutch frequency varied from 1 to 7 per breeding season, with female reproductive performance (determined by clutch frequency, annual fecundity, and annual reproductive output) maximized in females with a SVL of 68.0 mm. Accordingly to our hypothesis, the reproductive performance was maximized in the intermediate sized rather than the largest individuals in both sexes, and the body size maximizing reproductive performance was similar for both sexes. Future work could usefully investigate other lineages of lizards with sexually monomorphic species in a phylogenetic context to corroborate the hypothesis of this study.

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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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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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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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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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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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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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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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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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The present paper deals with the recovery growth experiment in Juvenile Grass Lizards, Takydromus septentrionalis, following short periods of fasting. Lizards were collected in April and May 2005 from a population in Chuzhou (Anhui, eastern China). The captured lizards were transported to our laboratory at Chuzhnu University, where they were maintained to in each 600 mm×450 mm×300 mm (length×width×height )glass terrarium, of which the bottom was filled with moist soil, grasses and debris. Lizards were kept starved at the nature temperature for 3d prior to feeding to insure post-absorptive conditions. 39 lizards were divided into 4 experimental groups, which were deprived of food for 0 (control), 3, 6 and 9 days, respectively. The experiment lasted for 24 days for each test group. Then each group was fed with larvae of Tenebrio molitor for the rest of the experimental period. During the recovery growth, wet mass in each test group increased with the lapsed time. Food intake in S3 and S9 were higher than in C. Apparent digestive coefficient (ADC) and assimilation efficiency (AE) in each group by starvation were similar than those in C. ADC and AE in S3 were higher than those in S6 and S9. It is suggested that: (1) T. septentrionalis starvation pretreatment for 3, 6, 9 days show significantly compensatory effect in the recovery growth; (2) the compensatory growth result from significant increase in feeding intensity in the recovery growth.

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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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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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The variation of water and energy contents in the Major Energy Reserves were studied in adult grass lizards (Takydromus septentrionalis) from March to September,2000,which collected month by month from Langyashan Mountain,Chuzhou,Anhui,Eastern China.Males and females differed in water contents of tail-free carcass and liver.There were significant monthly changes in water contents and energy contents of tail-free carcass,tail and liver in both males and females.Significant monthly changes in energy contents of fat bodies were also found in both sexes.Energy contents of tail-free carcass (including lean tail-free carcass),tail (including lean tail) were all higher in males than in females,but those of liver lower in males than in females.The results indicated that seasonal variation in the major energy reserves were correlated with the reproduction and winter hibernation.

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We acclimated adults of Takydromus septentrionalis (northern grass lizard) from four localities (populations) under identical thermal conditions to examine whether local thermal conditions have a fixed influence on thermal preference and thermal tolerance in the species. Selected body temperature (Tsel), critical thermal minimum (CTMin), and critical thermal maximum (CTMax) did not differ between sexes and among localities in lizards kept under identical laboratory conditions for approximately 5 months, and the interaction effects between sex and locality on these measures were not significant. Lizards acclimated to the three constant temperatures (20, 25, and 35 degrees C) differed in Tsel, CTMin, and CTMax. Tsel, CTMin, and CTMax all shifted upward as acclimation temperature increased, with Tsel shifting from 32.0 to 34.1 degrees C, CTMin from 4.9 to 8.0 degrees C, and CTMax from 42.0 to 44.5 degrees C at the change-over of acclimation temperature from 20 to 35 degrees C. Lizards acclimated to the three constant temperatures also differed in the range of viable body temperatures; the range was widest in the 25 degrees C treatment (38.1 degrees C) and narrowest in the 35 degrees C treatment (36.5 degrees C), with the 20 degrees C treatment in between (37.2 degrees C). The results of this study show that local thermal conditions do not have a fixed influence on thermal preference and thermal tolerance in T. septentrionalis.

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We acclimated adults of Takydromus septentrionalis (northern grass lizard) from four localities (populations) under identical thermal conditions to examine whether local thermal conditions have a fixed influence on thermal preference and thermal tolerance in the species. Selected body temperature (Tsel), critical thermal minimum (CTMin), and critical thermal maximum (CTMax) did not differ between sexes and among localities in lizards kept under identical laboratory conditions for approximately 5 months, and the interaction effects between sex and locality on these measures were not significant. Lizards acclimated to the three constant temperatures (20, 25, and 35 degrees C) differed in Tsel, CTMin, and CTMax. Tsel, CTMin, and CTMax all shifted upward as acclimation temperature increased, with Tsel shifting from 32.0 to 34.1 degrees C, CTMin from 4.9 to 8.0 degrees C, and CTMax from 42.0 to 44.5 degrees C at the change-over of acclimation temperature from 20 to 35 degrees C. Lizards acclimated to the three constant temperatures also differed in the range of viable body temperatures; the range was widest in the 25 degrees C treatment (38.1 degrees C) and narrowest in the 35 degrees C treatment (36.5 degrees C), with the 20 degrees C treatment in between (37.2 degrees C). The results of this study show that local thermal conditions do not have a fixed influence on thermal preference and thermal tolerance in T. septentrionalis.

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Adult grass lizards,Takydromus septentrionalis, are similar in size (SVL) but sexually dimorphic in head size (males larger than females ) throughout its range. A comprehensive analysis on the ecological and evolutionary sources of sexual differences in growth and size of heads requires a detailed understanding of growth trajectories during ontogeny.In this study,we specifically quesioned at what point during ontogeny males and females diverge in head size:at birth,during juvenile growth,or as mature adults? Our results indicate that males and females begin to diverge in head size at birth,although the divergence in the newly emerged young is much less pronounced than that in adults.An ANCOVA indicates that adult females even have smaller heads than do juveniles (including the newly emerged young).This suggests that adult females partition less resources into head growth but more into carcass growth,thereby leaving a larger space for eggs so as to increase reproductive output.Lizards collected in different seasons,from different populations,and at different ontogenetic stages,in various degrees,differ in food niche width and breadth.However,no direct evidence shows a substantial contribution of the divergence in head size to the segregation of food niche between males and females.Our analyses support that sexual selection is the main evolutionary source of smaller heads in female T.septentrionalis.

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We studied the ultrastructure of the spermatozoon of northern grass lizards (Takydromus septentrionalis,n=5) collected in April 2003 from a population in Wenzhou, Zhejiang, eastern China. The mature spermatozoa are characterized by: A circular acrosome; an acrosomal vesicle divided into cortex and medulla; the unilateral ridge of the acrosome vesicle divided into cortex and medulla, the electronlucent zone between them; a prenuclear perforatorium, absence of the perforatorium base plate; presence of the subacrosomal space; an elongated nucleus, absence of a endonuclear canal; absence of epinuclear lucent zone; rounded nuclear shoulders. A bilateral stratified laminar structure within the neck region. A short midpiece; absence of multilaminar membranes; in longitudinal section, two tiers of mitochondria, in transverse section, six mitochondria with linear cristae; the presence of two dense body groups, the ring structure consisting of the complete ring; the arrangement pattern of the ring structure and mitochondria in rs1/mi1 and rs2/mi2; a fibrous sheath in the midpiece; presence of an annulus. A thin granular zone of cytoplasm at the anterior portion of the principal piece; the enlarged fiber 3 and fiber 8 disappearing at the anterior portion of the principal piece; the axoneme complex having an usual 9+2 pattern. Within lacertid lizards, sperms differ among species in the number of mitochondria, but in all species studied so far, there are two groups of dense bodies. However, lizards of different taxa differ, in various degrees, in such sperm morphological traits as acrosome vesicle, subacrosomal space, epinuclear lucent zone, perforatorium base plate, nuclear shoulder and number and arrangement of mitochondria and dense bodies. These differences suggest that sperm morphology provides additional information which one may use to study the phylogeny of lizards.

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Animals in captivity undergo a range of environmental changes from wild animals. An increasing number of studies show that captivity significantly affects the abundance and community structure of gut microbiota. The northern grass lizard (Takydromus septentrionalis) is an extensively studied lacertid lizard and has a distributional range covering the central and southeastern parts of China. Nonetheless, little is known about the gut microbiota of this species, which may play a certain role in nutrient and energy metabolism as well as immune homeostasis. Here, we examined the differences in the gut microbiota between two groups (wild and captive) of lizards through 16S rRNA sequencing using the Illumina HiSeq platform. The results demonstrated that the dominant microbial components in both groups consisted of Proteobacteria, Firmicutes, and Tenericutes. The two groups did not differ in the abundance of these three phyla. Citrobacter was the most dominant genus in wild lizards, while Morganella was the most dominant genus in captive lizards. Moreover, gene function predictions showed that genes at the KEGG pathway levels2 were more abundant in wild lizards than in captive lizards but, at the KEGG pathway levels1, the differences in gene abundances between wild and captive lizards were not significant. In summary, captivity exerted a significant impact on the gut microbial community structure and diversity in T. septentrionalis, and future work could usefully investigate the causes of these changes using a comparative approach.