Length of activity season drives geographic variation in body size of a widely distributed lizard

Understanding the factors that drive geographic variation in life history is an important challenge in evolutionary ecology. Here, we analyze what predicts geographic variation in life‐history traits of the common lizard, Zootoca vivipara, which has the globally largest distribution range of all terrestrial reptile species. Variation in body size was predicted by differences in the length of activity season, while we found no effects of environmental temperature per se. Females experiencing relatively short activity season mature at a larger size and remain larger on average than females in populations with relatively long activity seasons. Interpopulation variation in fecundity was largely explained by mean body size of females and reproductive mode, with viviparous populations having larger clutch size than oviparous populations. Finally, body size‐fecundity relationship differs between viviparous and oviparous populations, with relatively lower reproductive investment for a given body size in oviparous populations. While the phylogenetic signal was weak overall, the patterns of variation showed spatial effects, perhaps reflecting genetic divergence or geographic variation in additional biotic and abiotic factors. Our findings emphasize that time constraints imposed by the environment rather than ambient temperature play a major role in shaping life histories in the common lizard. This might be attributed to the fact that lizards can attain their preferred body temperature via behavioral thermoregulation across different thermal environments. Length of activity season, defining the maximum time available for lizards to maintain optimal performance, is thus the main environmental factor constraining growth rate and annual rates of mortality. Our results suggest that this factor may partly explain variation in the extent to which different taxa follow ecogeographic rules.     

Climate and mammalian life histories

Mammals display considerable geographical variation in life history traits. To understand how climatic factors might influence this variation, we analysed the relationship between life history traits – adult body size, litter size, number of litters per year, gestation length, neonate body mass, weaning age and age at sexual maturity – and several environmental variables quantifying the seasonality and predictability of temperature and precipitation across the distribution range of five terrestrial mammal groups. Environmental factors correlated strongly with each other; therefore, we used principal components analysis to obtain orthogonal climatic predictors that could be used in multivariate models. We found that in bats, primates and even‐toed ungulates adult body size tends to be larger in species inhabiting cold, dry, seasonal environments, whereas in carnivores and rodents a smaller body size is characteristic of warm, dry environments, suggesting that low food availability might limit adult size. Species inhabiting cold, dry, seasonal habitats have fewer, larger litters and shorter gestation periods; however, annual fecundity in these species is not higher, implying that the large litter size of mammals living at high latitudes is probably a consequence of time constraints imposed by strong seasonality. On the other hand, the number of litters per year and annual fecundity were greater in species inhabiting environments with higher seasonality in precipitation. Lastly, we found little evidence for specific effects of environmental variability. Our results highlight the complex effects of environmental factors in the evolution of life history traits in mammals. © 2014 The Linnean Society of London, Biological Journal of the Linnean Society, 2014, 111 , 719–736.     

Macroclimatic and maternal effects on the evolution of reproductive traits in lizards

Much of life‐history theory rests on fundamental assumptions about constraints on the acquisition and allocation of energy to growth and reproduction. In general, the allocation of energy to reproduction depends on maternal size, which in turn depends on environmental factors experienced throughout the life of the mother. Here, we used phylogenetic path analyses to evaluate competing hypotheses about the environmental and maternal drivers of reproductive traits in lizards. In doing so, we discovered that precipitation, rather than temperature, has shaped the evolution of the life history. Specifically, environments with greater rainfall have enabled the evolution of larger maternal size. In turn, these larger mothers produce larger clutches of larger offspring. However, annual precipitation has a negative direct effect on offspring size, despite the positive indirect effect mediated by maternal size. Possibly, the evolution of offspring size was driven by the need to conserve water in dry environments, because small organisms are particularly sensitive to water loss. Since we found that body size variation among lizards is related to a combination of climatic factors, mainly precipitation and perhaps primary production, our study challenges previous generalizations (e.g., temperature‐size rule and Bergmann''s rule) and suggests alternative mechanisms underlying the evolution of body size.     

Life-history strategies of Australian lizards: a comparison between the tropics and the temperate zone

Summary Previous studies have suggested that tropical and temperate-zone lizards may differ fundamentally in life histories. We tested the applicability of this idea to Australian species by comparing temperate-zone species of agamid and scincid lizards with their congeners from the seasonal tropics. Data were derived from dissection of 1,941 specimens and from published information. Clutch size and egg size were positively correlated with mean maternal body size in most lizard species from both climatic zones. Mean body size of the lizards studies did not differ between the tropics and the temperate zone, nor did egg or hatchling size. However, tropical skinks showed considerably (approximately 20%) lower clutch size and relative clutch mass than did temperate-zone skinks. This difference was partly due to the higher incidence of species with low, invariant clutch size in the tropical lizard fauna (as seen in other continents as well), but primarily due to a trend for lineages (especially genera) with relatively high fecundity to be more common in the temperate zone than in the tropics. In contrast to studies on African lizards, our data suggested that modification of clutch size between areas has not occurred within genera: congeneric species from the tropics and temperate zone did not differ in clutch size. Production of more than one clutch per annum by individual females was common in both climatic zones. Tropical lizards may differ from temperate-zone species in showing higher reproductive frequencies, more rapid growth and earlier maturation. However, most of these effects may be due to phenotypic responses to environmental conditions (especially longer annual activity season), rather than to genetically based lifehistory adaptations.     

A comparative study of body size and clutch size across the parasitoid Hymenoptera

Across animal species, body size and clutch size often form part of a suite of associated life history traits, exemplified by the "fast-slow continuum" in mammals. Across the parasitoid Hymenoptera however, a major axis of life history variation is the development mode of the larva (koinobiosis versus idiobiosis), and body size and clutch size do not seem to form clear associations with this major axis. Here we use a large comparative data set and the latest phylogenetic information to explore hypotheses that might explain the variation in body size and clutch size across species in parasitoids. We find evidence for three novel evolutionary correlations: changes in the stage of host attacked by the parasitoid (i.e. egg, larva, pupa) significantly predict changes in both body size and clutch size, whilst in gregarious species changes to higher latitudes are associated with reduced clutch size. We also find a number of hypothesized cross-species (phenotypic) associations that, however, we cannot demonstrate are the result of evolutionary correlations: large bodied species in our data tend to lay small clutches; koinobionts are larger than idiobionts attacking the same host stage; tropical species are smaller than temperate species (Bergmann's rule). Our results provide support for theoretical models of trait evolution in parasitoids, whilst the associations between latitude and life history may help explain why species richness in the family Ichneumonidae peaks at intermediate latitudes. Our results also show the continuing value of phylogenetically-based comparative analyses and demonstrate that recent work on parasitoid phylogenetics has produced significant benefits for our understanding of life history evolution.     

Patterns and drivers of intraspecific variation in avian life history along elevational gradients: a meta‐analysis

Elevational gradients provide powerful natural systems for testing hypotheses regarding the role of environmental variation in the evolution of life‐history strategies. Case studies have revealed shifts towards slower life histories in organisms living at high elevations yet no synthetic analyses exist of elevational variation in life‐history traits for major vertebrate clades. We examined (i) how life‐history traits change with elevation in paired populations of bird species worldwide, and (ii) which biotic and abiotic factors drive elevational shifts in life history. Using three analytical methods, we found that fecundity declined at higher elevations due to smaller clutches and fewer reproductive attempts per year. By contrast, elevational differences in traits associated with parental investment or survival varied among studies. High‐elevation populations had shorter and later breeding seasons, but longer developmental periods implying that temporal constraints contribute to reduced fecundity. Analyses of clutch size data, the trait for which we had the largest number of population comparisons, indicated no evidence that phylogenetic history constrained species‐level plasticity in trait variation associated with elevational gradients. The magnitude of elevational shifts in life‐history traits were largely unrelated to geographic (altitude, latitude), intrinsic (body mass, migratory status), or habitat covariates. Meta‐population structure, methodological issues associated with estimating survival, or processes shaping range boundaries could potentially explain the nature of elevational shifts in life‐history traits evident in this data set. We identify a new risk factor for montane populations in changing climates: low fecundity will result in lower reproductive potential to recover from perturbations, especially as fewer than half of the species experienced higher survival at higher elevations.     

Life history patterns in lizards of the arid and semiarid zone of Australia

Summary Studies on the life histories and population dynamics of lizards in the semiarid/arid zone of Australia are reviewed to identify the influence of size (female mean snout-vent length), phylogeny (family effects) and ecological parameters on the evolution of life history traits of these species. Species producing more than one clutch per year are larger than single-clutched ones. In an ANCOVA, significant effects of size and phylogeny on clutch size and on age at sexual maturity were found. Microhabitat (arboreal, terrestrial, and subterranean life style) also had an effect on clutch size, but only mediated through a significant interaction with size. However, results of the ANCOVAs depend on the families and ecological parameters included in the analyses. Therefore, caution is necessary in interpreting or generalizing the results; in any case, size and phylogeny explain only a small percentage of the observed variation. Nevertheless, a direct comparison of a set of syntopic/paratopic desert lizards supports and extends the main conclusions of the ANCOVA. A significant but small phylogenetic effect was found, and arboreal microhabitat was associated with greater age at sexual maturity. Activity (diurnal versus nocturnal) influenced yearly mortalities and clutch frequencies. For both, microhabitat and activity, predation levels and size-dependent mortality were the likely selective factors causing these correlations. The demographic environment explains the paucity of duurnal lizard species with fixed clutch sizes in the semiarid/arid zone of Australia. Possible causes for the evolution of fixed clutch sizes are discussed.     

Evolution of biometric and life‐history traits in lizards (Gallotia) from the Canary Islands

The aim was to study as to how biometric and life‐history traits of endemic lacertids in the Canary Islands (genus Gallotia) may have evolved, and possible factors affecting the diversification process of this taxon on successively appearing islands have been deduced. To that end, comparative analyses of sexual dimorphism and scaling of different body, head and life‐history traits to body size in 10 species/subspecies of Gallotia have been carried out. Both Felsenstein's independent contrasts and Huey and Bennett's ‘minimum evolution’ analyses show that male and female snout‐vent length (SVL) changed proportionally (sexual size dimorphism not changing with body size) throughout the evolution of these lizards and all within‐sex biometric traits have changed proportionally to SVL. Life‐history traits (size at sexual maturity, clutch size, hatchling SVL and mass, and life span) are highly correlated with adult female body size, the first two being the only traits with a positive allometry to female SVL. These results, together with the finding that the slope of hatchling SVL to female SVL regression was lower than that of SVL at maturity to female SVL, indicates that larger females reach maturity at a larger size, have larger clutches and, at the same time, have relatively smaller hatchlings than smaller females. There was no significant correlation between any pair of life‐history traits after statistically removing the effect of body size. As most traits changed proportionally to SVL, the major evolutionary change has been that of body size (a ca. threefold change between the largest and the smallest species), that is suggested to be the effect of variable ecological conditions faced by founder lizards in each island.     

Geographic variation of life‐history traits in the sand lizard,Lacerta agilis: testing Darwin's fecundity‐advantage hypothesis

The fecundity‐advantage hypothesis (FAH) explains larger female size relative to male size as a correlated response to fecundity selection. We explored FAH by investigating geographic variation in female reproductive output and its relation to sexual size dimorphism (SSD) in Lacerta agilis, an oviparous lizard occupying a major part of temperate Eurasia. We analysed how sex‐specific body size and SSD are associated with two putative indicators of fecundity selection intensity (clutch size and the slope of the clutch size–female size relationship) and with two climatic variables throughout the species range and across two widespread evolutionary lineages. Variation within the lineages provides no support for FAH. In contrast, the divergence between the lineages is in line with FAH: the lineage with consistently female‐biased SSD (L. a. agilis) exhibits higher clutch size and steeper fecundity slope than the lineage with an inconsistent and variable SSD (L. a. exigua). L. a. agilis shows lower offspring size (egg mass, hatchling mass) and higher clutch mass relative to female mass than L. a. exigua, that is both possible ways to enhance offspring number are exerted. As the SSD difference is due to male size (smaller males in L. a. agilis), fecundity selection favouring larger females, together with viability selection for smaller size in both sexes, would explain the female‐biased SSD and reproductive characteristics of L. a. agilis. The pattern of intraspecific life‐history divergence in L. agilis is strikingly similar to that between oviparous and viviparous populations of a related species Zootoca vivipara. Evolutionary implications of this parallelism are discussed.     

Allometry of reproduction in two species of gekkonid lizards (Gehyra): effects of body size miniaturization on clutch and egg sizes

In squamate reptiles there is an allometric pattern for small-bodied females to have smaller clutches and proportionally larger eggs than large-bodied females, and this pattern occurs both among and within species. The allometric patterns in two species of the gecko Gehyra were studied to see how evolutionary reductions in adult body size affect fecundity and offspring size among species, and how these changes affect allometric relationships within species. Gehyra dubia has two eggs per clutch (the typical clutch size for gekkonid lizards), whereas the smallerbodied G. variegata has a single egg per clutch. Within both species, egg size increased with female body size. The data are consistent with at least two mechanistic hypotheses: (1) that the width of the pelvis constrains egg size; and (2) in species with invariant clutch sizes, larger females can only allocate additional energy towards egg size and not number. More direct tests of these hypotheses are warranted. Miniaturization of body sizes in Gehyra is correlated with a clutch size reduction of 50% (from two to one), and a large (1.7-fold) compensatory increase in relative egg mass. However, the small-bodied G. variegata (one egg per clutch) had a lower relative clutch mass than did G. dubia. These findings have implications for understanding the influence of evolutionary reductions in body size on reproductive traits, and for allometric trends in squamate reptiles in general.     

Global warming and positive fitness response in mountain populations of common lizards Lacerta vivipara

Recent global warming threatens many species and has already caused population‐ and species‐level extinctions. In particular, high risks of extinction are expected for isolated populations of species with low dispersal abilities. These predictions rely on widely used ‘climatic envelope’ models, while individual responses, the ultimate driver of a species response to climate change, have been most often neglected. Here, we report on some changes in life‐history traits of a dispersal‐limited reptile species (a poorly studied taxa) living in isolated populations. Using long‐term data on common lizards collected in southern France, we show that individual body size dramatically increased in all the four populations studied over the past 18 years. This increase in body size in all age classes appeared related to a concomitant increase in temperature experienced during the first month of life (August). Daily maximum temperature in August increased by 2.2°C and yearling snout‐vent‐length increased by about 28%. As a result, adult female body size increased markedly, and, as fecundity is strongly dependent on female body size, clutch size and total reproductive output also increased. For one population where capture–recapture data were available, adult survival was positively related to May temperature. All fitness components investigated therefore responded positively to the increase in temperature, such that it might be concluded that the common lizard has been advantaged by the shift in temperature. We contrast these short‐term results with the long‐term habitat‐based prediction that these populations located close to mountain tops on the southern margin of the species range should be unable to cope with the alteration of their habitat. To achieve a better prediction of a species persistence, one will probably need to combine both habitat and individual‐based approaches.     

A comparative analysis of dispersal syndromes in terrestrial and semi‐terrestrial animals

Dispersal, the behaviour ensuring gene flow, tends to covary with a number of morphological, ecological and behavioural traits. While species‐specific dispersal behaviours are the product of each species’ unique evolutionary history, there may be distinct interspecific patterns of covariation between dispersal and other traits (‘dispersal syndromes’) due to their shared evolutionary history or shared environments. Using dispersal, phylogeny and trait data for 15 terrestrial and semi‐terrestrial animal Orders (> 700 species), we tested for the existence and consistency of dispersal syndromes across species. At this taxonomic scale, dispersal increased linearly with body size in omnivores, but decreased above a critical length in herbivores and carnivores. Species life history and ecology significantly influenced patterns of covariation, with higher phylogenetic signal of dispersal in aerial dispersers compared with ground dwellers and stronger evidence for dispersal syndromes in aerial dispersers and ectotherms, compared with ground dwellers and endotherms. Our results highlight the complex role of dispersal in the evolution of species life‐history strategies: good dispersal ability was consistently associated with high fecundity and survival, and in aerial dispersers it was associated with early maturation. We discuss the consequences of these findings for species evolution and range shifts in response to future climate change.     

Pleistocene climatic changes drive diversification across a tropical savanna

Spatial responses of species to past climate change depend on both intrinsic traits (climatic niche breadth, dispersal rates) and the scale of climatic fluctuations across the landscape. New capabilities in generating and analysing population genomic data, along with spatial modelling, have unleashed our capacity to infer how past climate changes have shaped populations, and by extension, complex communities. Combining these approaches, we uncover lineage diversity across four codistributed lizards from the Australian Monsoonal Tropics and explore how varying climatic tolerances interact with regional climate history to generate common vs. disparate responses to late Pleistocene change. We find more divergent spatial structuring and temporal demographic responses in the drier Kimberley region compared to the more mesic and consistently suitable Top End. We hypothesize that, in general, the effects of species’ traits on sensitivity to climate fluctuation will be more evident in climatically marginal regions. If true, this points to the need in climatically marginal areas to craft more species‐(or trait)‐specific strategies for persistence under future climate change.     

Sexual dimorphism of heads and abdomens: Different approaches to ‘being large’ in female and male lizards

Sexual‐size dimorphism (SSD) is widespread in animals. Body length is the most common trait used in the study of SSD in reptiles. However, body length combines lengths of different body parts, notably heads and abdomens. Focusing on body length ignores possible differential selection pressures on such body parts. We collected the head and abdomen lengths of 610 lizard species (Reptilia: Squamata: Sauria). Across species, males have relatively larger heads, whereas females have relatively larger abdomens. This consistent difference points to body length being an imperfect measure of lizard SSD because it comprises both abdomen and head lengths, which often differ between the sexes. We infer that female lizards of many species are under fecundity selection to increase abdomen size, consequently enhancing their reproductive output (enlarging either clutch or offspring size). In support of this, abdomens of lizards laying large clutches are longer than those of lizards with small clutches. In some analyses, viviparous lizards have longer abdomens than oviparous lizards with similar head lengths. Our data also suggest that male lizards are under sexual selection to increase head size, which is positively related to winning male–male combats and to faster grasping of females. Thus, larger heads could translate into higher probability to mate. © 2013 The Linnean Society of London, Biological Journal of the Linnean Society, 2013, 110 , 665–673.     

The interplay between natural and sexual selection in the evolution of sexual size dimorphism in Sceloporus lizards (Squamata: Phrynosomatidae)

Sexual size dimorphism (SSD) evolves because body size is usually related to reproductive success through different pathways in females and males. Female body size is strongly correlated with fecundity, while in males, body size is correlated with mating success. In many lizard species, males are larger than females, whereas in others, females are the larger sex, suggesting that selection on fecundity has been stronger than sexual selection on males. As placental development or egg retention requires more space within the abdominal cavity, it has been suggested that females of viviparous lizards have larger abdomens or body size than their oviparous relatives. Thus, it would be expected that females of viviparous species attain larger sizes than their oviparous relatives, generating more biased patterns of SSD. We test these predictions using lizards of the genus Sceloporus. After controlling for phylogenetic effects, our results confirm a strong relationship between female body size and fecundity, suggesting that selection for higher fecundity has had a main role in the evolution of female body size. However, oviparous and viviparous females exhibit similar sizes and allometric relationships. Even though there is a strong effect of body size on female fecundity, once phylogenetic effects are considered, we find that the slope of male on female body size is significantly larger than one, providing evidence of greater evolutionary divergence of male body size. These results suggest that the relative impact of sexual selection acting on males has been stronger than fecundity selection acting on females within Sceloporus lizards.     

宁波滑蜥两性异形和雌性繁殖

蜥蜴的雌性繁殖特征对理解两性异形的进化原因起着重要作用。于2011年4月在安徽滁州采集宁波滑蜥(Scincella modesta),定量研究该种形态特征的两性异形和雌性繁殖特征,检验成体形态特征两性异形与雌性繁殖的相关性。研究共采集43条(17♀♀,26♂♂)宁波滑蜥,雄性和雌性个体的最大体长分别为47.4 mm和46.6 mm。雌雄两性在体长和头宽上没有差异,而在腹长和头长上差异显著,雄性有较大的头长,雌性有较大的腹长。宁波滑蜥年产单窝卵。窝卵数和窝卵重与雌体体长及腹长呈正相关,卵重与雌体体长无相关性。窝卵数及卵重的变异系数分别为0.20和0.12。卵长径与窝卵数呈负相关,而卵短径与窝卵数无关。雌体主要通过增加窝卵数来增加繁殖输出。这些结果表明,宁波滑蜥是雌雄个体大小同形的两性异形模式,性选择使得雄性有着较大的头长,以具有较高的交配成功率,生育力选择使得雌性有着较大的腹长,以具有较大的生育力和繁殖输出。     

Interspecific competition affects evolutionary links between cavity nesting,migration and clutch size in Old World flycatchers (Muscicapdae)

The ecology of cavity nesting in passerine birds has been studied extensively, yet there are no phylogenetic comparative studies that quantify differences in life history traits between cavity‐ and open‐nesting birds within a passerine family. We test existing hypotheses regarding the evolutionary significance of cavity nesting in the Old World flycatchers (Muscicapidae). We used a multi‐locus phylogeny of 252 species to reconstruct the evolutionary history of cavity nesting and to quantify correlations between nest types and life history traits. Within a phylogenetic generalized linear model framework, we found that cavity‐nesting species are larger than open‐nesting species and that maximum clutch sizes are larger in cavity‐nesting lineages. In addition to differences in life history traits between nest types, species that breed at higher latitudes have larger average and maximum clutch sizes and begin to breed later in the year. Gains and losses of migratory behaviour have occurred far more often in cavity‐nesting lineages than in open‐nesting taxa, suggesting that cavity nesting may have played a crucial role in the evolution of migratory behaviour. These findings identify important macro‐evolutionary links between the evolution of cavity nesting, clutch size, interspecific competition and migratory behaviour in a large clade of Old World songbirds.     

Rain,prey and predators: climatically driven shifts in frog abundance modify reproductive allometry in a tropical snake

To predict the impacts of climate change on animal populations, we need long-term data sets on the effects of annual climatic variation on the demographic traits (growth, survival, reproductive output) that determine population viability. One frequent complication is that fecundity also depends upon maternal body size, a trait that often spans a wide range within a single population. During an eight-year field study, we measured annual variation in weather conditions, frog abundance and snake reproduction on a floodplain in the Australian wet-dry tropics. Frog numbers varied considerably from year to year, and were highest in years with hotter wetter conditions during the monsoonal season (“wet season”). Mean maternal body sizes, egg sizes and post-partum maternal body conditions of frog-eating snakes (keelback, Tropidonophis mairii, Colubridae) showed no significant annual variation over this period, but mean clutch sizes were higher in years with higher prey abundance. Larger females were more sensitive to frog abundance in this respect than were smaller conspecifics, so that the rate at which fecundity increased with body size varied among years, and was highest when prey availability was greatest. Thus, the link between female body size and reproductive output varied among years, with climatic factors modifying the relative reproductive rates of larger (older) versus smaller (younger) animals within the keelback population.     

Latitudinal and climatic variation in body size and dorsal scale counts in Sceloporus lizards:a phylogenetic perspective

Squamates often follow an inverse Bergmann's rule, with larger-bodied animals occurring in warmer areas or at lower latitudes. The size of dorsal scales in lizards has also been proposed to vary along climatic gradients, with species in warmer areas exhibiting larger scales, putatively to reduce heat load. We tested for these patterns in the diverse and widespread lizard genus Sceloporus. Among 106 species or populations, body size was associated positively with maximum temperature (consistent with the inverse of Bergmann's rule) and aridity, but did not covary with latitude. Scale size (inferred from the inverse relation with numbers of scales) was positively related to body size. Controlling for body size via multiple regression, scale size was associated negatively with latitude (best predictor), positively with minimum temperature, and negatively with aridity (similar results were obtained using scores from a principal components analysis of latitude and climatic indicators). Thus, lizards with larger scales are not necessarily found in areas with higher temperatures. Univariate analyses indicated phylogenetic signal for body size, scale counts, latitude, and all climate indicators. In all cases, phylogenetic regression models fit the data significantly better than nonphylogenetic models; thus, residuals for log(10) number of dorsal scale rows exhibited phylogenetic signal.     

Identifying sources of variation in reproductive and life-history traits among five populations of a Chinese lizard (Takydromus septentrionalis, Lacertidae)

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 maintenance 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 offspring size to fecundity.  © 2005 The Linnean Society of London, Biological Journal of the Linnean Society , 2005, 85 , 443–453.