山地麻蜥的雌性繁殖和孵出幼体特征

山地麻蜥年产多窝卵。窝卵数、窝卵重和卵重与雌体体长呈正相关,卵重与窝卵数无关。窝卵数、窝卵重和卵重不存在明显的季节变化。卵长径与卵短径呈正相关,卵长径、卵短径与窝卵数无关。雌体通过增加窝卵数和卵大小增加繁殖输出。孵出幼体的体重、体长、尾长、躯干干重、刺余卵黄不存在季节差异,但5月份孵出幼体的脂肪体高于4月份。     

Insular shifts and trade-offs in life-history traits in pond frogs in the Zhoushan Archipelago, China

Island and mainland populations of animal species often differ strikingly in life-history traits such as clutch size, egg size, total reproductive effort and body size. However, despite widespread recognition of insular shifts in these life-history traits in birds, mammals and reptiles, there have been no reports of such life-history shifts in amphibians. Furthermore, most studies have focused on one specific life-history trait without explicit consideration of coordinated evolution among these intimately linked life-history traits, and thus the relationships among these traits are poorly studied. Here we provide the first evidence of insular shifts and trade-offs in a coordinated suite of life-history traits for an amphibian species, the pond frog Rana nigromaculata . Life-history data were collected from eight islands in the Zhoushan Archipelago and neighboring mainland China. We found consistent, significant shifts in all life-history traits between mainland and island populations. Island populations had smaller clutch sizes, larger egg sizes, larger female body size and invested less in total reproductive effort than mainland populations. Significant negative relationships were found between egg size and clutch size and between egg size and total reproductive effort among frog populations after controlling for the effects of body size. Therefore, decreased reproductive effort and clutch size, larger egg size and body size in pond frogs on islands were selected through trade-offs as an overall life-history strategy. Our findings contribute to the formation of a broad, repeatable ecological generality for insular shifts in life-history traits across a range of terrestrial vertebrate taxa.     

Measuring temporal variation in reproductive output reveals optimal resource allocation to reproduction in the northern grass lizard, Takydromus septentrionalis

We measured the reproductive output of Takydromus septentrionalis collected over 5 years between 1997 and 2005 to test the hypothesis that reproductive females should allocate an optimal fraction of accessible resources in a particular clutch and to individual eggs. Females laid 1–7 clutches per breeding season, with large females producing more, as well as larger clutches, than did small females. Clutch size, clutch mass, annual fecundity, and annual reproductive output were all positively related to female size (snout–vent length). Females switched from producing more, but smaller eggs in the first clutch to fewer, but larger eggs in the subsequent clutches. The mass-specific clutch mass was greater in the first clutch than in the subsequent clutches, but it did not differ among the subsequent clutches. Post-oviposition body mass, clutch size, and egg size showed differing degrees of annual variation, but clutch mass of either the first or the second clutch remained unchanged across the sampling years. The regression line describing the size–number trade-off was higher in the subsequent clutch than in the first clutch, but neither the line for first clutch, nor the line for the second clutch varied among years. Reproduction retarded growth more markedly in small females than in large ones. Our data show that: (1) trade-offs between size and number of eggs and between reproduction and growth (and thus, future reproduction) are evident in T. septentrionalis ; (2) females allocate an optimal fraction of accessible resources in current reproduction and to individual eggs; and (3) seasonal shifts in reproductive output and egg size are determined ultimately by natural selection.  © 2007 The Linnean Society of London, Biological Journal of the Linnean Society , 2007, 91 , 315–324.     

Life-history variation within a three-spined stickleback population in the Camargue

Among individuals of female three-spined sticklebacks Gasterosteus aculeatus from a population in the Camargue, southern France, studied in 12 successive years, adult L _T ranged from 31–64 mm, clutch size ranged from 33–660 eggs, and mean egg diameter per clutch ranged from 1.15–1.67 mm. Because the population was strictly annual, inter-annual variation corresponded to variation among generations having experienced different environmental conditions. Body mass varied significantly among years, suggesting an effect of varying environmental conditions. Gonad mass and clutch size increased with body mass, but mean egg diameter was not correlated to body mass. Body mass-adjusted gonad mass, interpreted as reproductive effort per clutch, did not vary significantly among years, suggesting that this trait was not influenced by environmental conditions. Body mass-adjusted clutch size and egg size varied significantly among years. Inter-annual variation in body length at breeding, clutch size and egg size was of the same order of magnitude as inter-population variation reported by other authors for this species. During the breeding season, reproductive effort and clutch size tended to increase. Egg size tended to decrease during the breeding season but this seasonal pattern varied among years. Observed life-history variation is discussed both in terms of its evolutionary significance and methodological implications in the study of life-history variation among populations.     

中国石龙子雌体繁殖特征和卵孵化的地理变异

浙江丽水和广东韶关中国石龙子均年产单窝卵,窝卵数,窝卵重和卵重均与雌体SVL呈正相关,雌体头部形态,繁殖特征,产卵起始时间和孵孵化的热依赖性等有显著的地理变异;韶关石龙子产卵起始时间为5月中旬,比丽水经子约早两周,韶关石龙子窝卵数较大,卵较小,窝卵重与丽水石龙子无显著差异。韶关石龙子特定SVL的窝卵数比丽水石龙子多2．8枚卵,中国经子卵数量和大小之间有种群间权衡,无种数内权衡,同一种群内卵数量与卵大小无关,孵化温度影响石龙子孵出幼体的一些特征,24℃孵出细幼体比32℃孵出幼体大,躯干发育好,剩余卵黄少,韶关24℃孵出幼体的体重,躯干干重小于丽水幼体,韶关32℃孵出幼体的SVL小于丽水幼体,剩余卵黄大于丽水幼体,表明适宜卵孵化温度范围有地理变异。丽水石龙子卵对极端高温和低温的耐受性较强,适宜卵孵化温度范围较宽。     

Maternal body-volume as a constraint on reproductive output in lizards: evidence from the evolution of viviparity

A few species of squamate reptiles contain both oviparous (egg-laying) and viviparous (live-bearing) populations, and thus offer exceptional opportunities to test adaptationist hypotheses on the determinants of reproductive output. We focus on the hypothesis that maternal body-volume constrains reproductive output in squamate reptiles. If females are “full” of eggs, what happens when viviparity evolves within a lineage? Eggs increase in volume and mass during development, primarily due to the uptake of water, so how can they be accommodated within the mother's abdomen? We predict that the resultant increase in relative clutch mass (RCM) will be lessened by (1) a decrease in reproductive output (by reducing the number or size of offspring), and/or (2) an increase in maternal body-volume (via modifications of size or shape of adult females). Our comparisons of conspecific oviparous and viviparous lizards (Lerista bougainvillii) confirm that live-bearers carry heavier clutches (in both absolute and relative terms) and show the predicted shifts in body size and shape of reproductive females. However, offspring size and number were unaffected by the evolution of viviparity, and the shifts in maternal morphology were too small to fully offset the increase in clutch mass. Thus, RCMs increased by 50%, indicating that viviparous females produced clutches which more completely filled the space available in the abdominal cavity. We conclude that maternal body-volume does play a role in determining reproductive output, but that the observed clutch masses may be optimized, rather than maximized, with respect to the abdominal space available.     

Geographic variation in 'costs of reproduction' in the scincid lizard Lampropholis guichenoti

1. A major focus of life-history research has been the analysis of reproductive effort (RE). However, while clearly defined in theory, RE has proved very difficult to measure. Consequently, researchers have looked for indices that estimate the components of RE. Uncritical use of indices of reproductive investment, such as measures of costs of reproduction, without verification of the underlying assumptions of the link between these traits, may lead to spurious conclusions. In this paper, the common assumption that the physical burden of the clutch impairs locomotor ability in gravid reptiles is examined.
2. Two neighbouring populations of Australian scincid lizards ( Lampropholis guichenoti) are similar in adult body sizes, body shapes and reproductive output (egg sizes, clutch sizes, relative clutch masses).
3. Despite morphological and reproductive similarities, the effects of pregnancy on maternal locomotor ability (running speeds, as measured in a laboratory raceway) differed dramatically between the two populations.
4. Lizards from the two populations ran at similar speeds when nongravid (i.e. after egg-laying), but pregnancy significantly reduced running speeds in one population and increased them in the other. Thus, superficial similarities in body size and reproductive output masked a strong divergence in the locomotor 'costs' of reproduction.
5. Caution is advised if using simple measures of reproductive output (e.g. relative clutch mass) as indices of reproductive effort, or to generalize results even among conspecific populations.     

Food habits, growth rates, and reproductive biology of grass snakes, Natrix natrix (Colubridae) in the Italian Alps

A five-year mark-recapture study at Sella Nevea, a montane (1100 m a.s.1.) site in the Carnic Alps, provided information on diets, growth rates, and reproductive output in an Italian population of the wide-ranging grass snake, Natrix natrix. Our snakes resembled a previously-studied population in lowland Sweden in terms of body size at sexual maturation in females (70 cm) and mean adult female body length (82 cm). However, growth rates were lower in our population, and sexual maturation was delayed (6–8 years, versus 4–5 years in Sweden), perhaps because of the cool climate and relatively brief growing period each year. Females produced a single clutch of 4–24 eggs in late July each year. Larger females produced larger clutches, but clutch size relative to maternal size was lower than in Swedish grass snakes. Hatchling sizes and Relative Clutch Masses (RCMs) did not shift with increasing female size. RCMs may provide a useful index of 'costs of reproduction' in this population, because females with high RCMs were very emaciated after oviposition, and hence may experience a greater risk of mortality, as well as a high energy expenditure. Prolonged incubation gave rise to longer, thinner hatchlings, but the low environmental temperatures at the study site may favour early hatching (and hence, result in a shorter fatter hatchling emerging from the egg, with more of its energy stores unused). Compared to sympatric viviparous snakes ( Coronella austriaca and Vipera berus ), the oviparous grass snakes can achieve a much higher reproductive output owing to a larger clutch size and more frequent reproduction (annual, rather than biennial or triennial). The abundant prey resource used by grass snakes (amphibians) may also enable them to recoup energy more rapidly after reproduction; dietary composition shifts ontogenetically in both sexes, with the largest prey (mice and adult toads) taken primarily by large female snakes.     

Phenotypic plasticity in reproductive traits induced by food availability in a lacertid lizard, Takydromus septentrionalis

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. septentrionali s "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.     

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.     

蜡皮蜥的两性异形和繁殖输出

为研究蜡皮蜥(Leiolepis reevesii)两性异形和繁殖输出,于2002、2003年4月下旬从海南乐东一种群捕获423头蜡皮蜥。经检测得到繁殖雌体的最小体长为89．0mm,据此判定≥89．0mm的个体为性成熟。研究结果表明：①蜡皮蜥具有两性异形,雄性大于雌性且具有较大的头部。成体雄性头长和头宽随体长的增长速率大于雌性,幼体头长和头宽随体长的增长速率无显著的两性差异。以性别和年龄(成、幼体)为因子的双因子ANOVA比较两性头长和头宽与体长的回归剩余值发现,雄性头部大于雌性,幼体头部相对大于成体。②饲养于实验室的母体中有42头于2002、2003年5月22日～7月16日产出正常卵,这些繁殖雌体具有年产多窝卵的潜力。窝卵数和卵重的变异系数分别为0．18和0．13,前者变异度大于后者。窝卵数、窝卵重和卵重均与母体体长无关。卵重与相对生育力之间无显著的负相关性,表明蜡皮蜥缺乏卵数量与卵大小之间的权衡。相对窝卵重与母体体长呈显著的负相关,表明较小的母体具有相对较大的繁殖输出。因雌体繁殖会滞缓其生长,小母体具有相对较大的繁殖输出,至少部分地解释了雌性蜡皮蜥的成体为什么个体较小。     

Variation and trade-off in reproductive output of the Andean frog<Emphasis Type="Italic"> Hyla labialis</Emphasis>

I analysed variation in number and size of eggs in association with variation in body size of Hyla labialis females from three altitudinally widely separated populations in the Colombian Andes. Females at 3,500 m were significantly larger and heavier than those at 2,700 m, which in turn were larger and heavier than those at 2,000 m. After accounting for interpopulational differences in body size, clutch size (i.e. number of eggs) and egg size differed significantly between populations, but clutch mass did not. At 3,500 m, large relative clutch mass tended to be associated with larger eggs, whereas at lower altitudes it was associated with relatively larger clutch size. Within each population, females showed a significant trade-off between clutch size and egg size, even though relative clutch mass varied substantially and a clutch rarely used up the carrying capacity of their body cavity. At 3,500 m, comparisons between the early and late breeding season within a year suggested that climate fluctuations may play a role in reproductive-output variation within a population. Laboratory-kept females in a high somatic condition after spawning laid larger clutches in the next breeding event, but the trade-off persisted despite ad libitum food supply. Comparisons between field and laboratory suggest that limits to reproductive output were due to physiological, but not morphological restrictions. Therefore, the trade-off is better explained by energy limitation for egg growth depending on clutch size, than by the body cavity limitation hypothesis.     

Microgeographical variation in brown trout reproductive traits: possible effects of biotic interactions

This study documents substantial variation in reproductive traits among populations of stream-dwelling brown trout ( Salmo trutta L.) at a very small geographic scale. Within two streams, we found a parallel pattern of variation, where females living above major waterfalls produced fewer and larger eggs than conspecifics from below the waterfalls. Four additional streams were represented with either a below-waterfall site ( n =2) or an above-waterfall site ( n =2). When these streams were included in the analyses, there was no consistent difference in reproductive traits between females from above- and below-waterfall sites. There was no significant difference in total reproductive investment among sites within streams, but considerable variation among streams. Female first-year growth rates was estimated from scales, and differed significantly among populations. Within streams, females from below waterfalls experienced higher first-year growth rates as compared to females from above the waterfalls. Within seven out of eight populations, egg size increased significantly with increasing female body length. Within three populations, we found evidence for a trade-off between offspring size and offspring number, as a negative association between fecundity and egg size independently of adult body size. Within three populations egg size decreased significantly with increasing maternal first-year growth, independently of adult body size. We suggest that the within-stream differences in offspring size/number strategies are influenced by population density and growth effects. Earlier, we have shown that population densities are consistently lower below the waterfalls in these streams. The Alpine bullhead ( Cottus poecilopus ) is found only below the waterfalls and could influence brown trout demography.     

The use of ultrasonography to assess reproductive investment and output in pythons

Reproductive investment and output are integral fitness components, often incorporated into life‐history trade‐off models and important to population dynamics. The trade‐offs associated with reproduction can be dramatic in species such as snakes that make especially large investments into reproduction. Unfortunately, traditional methods used to determine reproductive investment and output are effective in many (but not all) situations. Thus, we used portable ultrasonography to serially estimate reproductive investment and reproductive output in three python species that exhibit significant variation in phylogeny, geographic range, body size, egg size, and clutch size: ball pythons (Python regius), Children's pythons (Antaresia childreni), and water pythons (Liasis fuscus). At each time point of measurement (range: 1–49 days pre‐oviposition), ultrasound estimates of viable clutch size were highly accurate in all three species. However, ultrasound estimates of mean viable egg mass, and thus viable clutch mass, significantly differed from the actual values (range: 23–73% error). Interestingly, this error was considerably smaller as females approached oviposition, suggesting that female pythons transfer a significant amount of water into their eggs during the week before oviposition. Thus, water balance during late‐stage egg development may be an integral part of reproductive success. The results obtained in the present study form the foundation for future assessments of reproductive investment, and also provide insight into the use of ultrasound technology to assist such efforts. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 772–778.     

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.     

A consistent long-lasting pattern of spatial variation in egg size and shape in blue tits (<Emphasis Type="Italic">Cyanistes caeruleus</Emphasis>)

Background

Interspecies variation in avian egg shape and size is understandable in terms of adaptation, allometry and phylogeny. Within-species variation in egg properties influences offspring fitness and can be explained by differences in allocation of resources into reproductive components of life history in mulidimensionally variable environments. Egg size is inherently traded-off with clutch size, which may also be true of egg shape in some cases. We investigated long-term variation in egg shape and size between two geographically close populations of blue tits Cyanistes caeruleus in relation to clutch size and habitat differences.

Results

The main finding is that there exists a persistent long-lasting pattern of spatial variation of egg size and shape between the two study populations of blue tits, 10 km apart, controlling for clutch size. Eggs in the urban park site were on average larger in volume and less spherical in shape than eggs in the forest site over 12 years of this study. Egg sizes were negatively associated with clutch sizes. Egg shape was not correlated with clutch size.

Conclusions

Our findings suggest that the pattern of variation in egg size and shape results from different trophic richness of the breeding habitats of the study populations, demanding different allocation of resources and, especially, from the contrasting difference in the availability of calcium.

     

Understanding reproductive allometry in turtles: A slippery “slope”

Measures of reproductive output in turtles are generally positively correlated with female body size. However, a full understanding of reproductive allometry in turtles requires logarithmic transformation of reproductive and body size variables prior to regression analyses. This allows for slope comparisons with expected linear or cubic relationships for linear to linear and linear to volumetric variables, respectively. We compiled scaling data using this approach from published and unpublished turtle studies (46 populations of 25 species from eight families) to quantify patterns among taxa. Our results suggest that for log–log comparisons of clutch size, egg width, egg mass, clutch mass, and pelvic aperture width to shell length, all scale hypoallometrically despite theoretical predictions of isometry. Clutch size generally scaled at ~1.7 to 2.0 (compared to an isometric expectation of 3.0), egg width at ~0.5 (compared to an expectation of 1.0), egg mass at ~1.1 to 1.3 (3.0), clutch mass at ~2.5 to 2.8 (3.0), and pelvic aperture width at 0.8–0.9 (1.0). We also found preliminary evidence that scaling may differ across years and clutches even in the same population, as well as across populations of the same species. Future investigators should aspire to collect data on all these reproductive parameters and to report log–log allometric analyses to test our preliminary conclusions regarding reproductive allometry in turtles.     

Not putting all their eggs in one basket: bet‐hedging despite extraordinary annual reproductive output of desert tortoises

Bet‐hedging theory makes the counter‐intuitive prediction that, if juvenile survival is low and unpredictable, organisms should consistently reduce short‐term reproductive output to minimize the risk of reproductive failure in the long‐term. We investigated the long‐term reproductive output of an Agassiz's desert tortoise (Gopherus agassizii) population and conformance to a bet‐hedging strategy of reproduction in an unpredictable but comparatively productive environment. Most females reproduced every year, even during periods of low precipitation and poor germination of food plants, and the mean percentage of reproducing females did not differ significantly on an annual basis. Although mean annual egg production (clutch size × clutch frequency) differed significantly among years, mean clutch size and mean clutch frequency remained relatively constant. During an El Niño year, mean annual egg production and mean annual clutch frequency were the highest ever reported for this species. Annual egg production was positively influenced by maternal body size but clutch size and clutch frequency were not. Our long‐term results confirm earlier conclusions based on short‐term research that desert tortoises have a bet‐hedging strategy of producing small clutches almost every year. The risk of long‐term reproductive failure is minimized in unpredictable environments, both through time by annually producing multiple small clutches over a long reproductive lifespan, even in years of low resource availability, and through space by depositing multiple annual clutches in different locations. The extraordinary annual reproductive output of this population appears to be the result of a typically high but unpredictable biomass of annual food plants at the site relative to tortoise habitat in dryer regions. Under the comparatively productive but unpredictable conditions, tortoises conform to predictions of a bet‐hedging strategy of reproduction with relatively small but consistent clutch sizes. Published 2015. This article is a U.S. Government work and is in the public domain in the USA, Biological Journal of the Linnean Society, 2015, 115 , 399–410.     

Bony pelvic canal size and shape in relation to body proportionality in humans

Obstetric selection acts on the female pelvic canal to accommodate the human neonate and contributes to pelvic sexual dimorphism. There is a complex relationship between selection for obstetric sufficiency and for overall body size in humans. The relationship between selective pressures may differ among populations of different body sizes and proportions, as pelvic canal dimensions vary among populations. Size and shape of the pelvic canal in relation to body size and shape were examined using nine skeletal samples (total female n = 57; male n = 84) from diverse geographical regions. Pelvic, vertebral, and lower limb bone measurements were collected. Principal component analyses demonstrate pelvic canal size and shape differences among the samples. Male multivariate variance in pelvic shape is greater than female variance for North and South Africans. High‐latitude samples have larger and broader bodies, and pelvic canals of larger size and, among females, relatively broader medio‐lateral dimensions relative to low‐latitude samples, which tend to display relatively expanded inlet antero‐posterior (A‐P) and posterior canal dimensions. Differences in canal shape exist among samples that are not associated with latitude or body size, suggesting independence of some canal shape characteristics from body size and shape. The South Africans are distinctive with very narrow bodies and small pelvic inlets relative to an elongated lower canal in A‐P and posterior lengths. Variation in pelvic canal geometry among populations is consistent with a high degree of evolvability in the human pelvis. Am J Phys Anthropol 151:88–101, 2013. © 2013 Wiley Periodicals, Inc.     

Variation of reproductive traits in a population of the lizard Lacerta vivipara

Variation in reproductive traits (sexual maturity, clutch size, clutch weight, mean egg mass, newborn weight) was studied during a four year period in a population of the live-bearing lizard Lacerta vivipara . Sexual maturity was associated with attaining a minimum body size. Clutch size increased with female body length and litter weight increased with clutch size. A major component of the within year variation in these reproductive traits was attributable to female size. Analysis of successive clutches in individual females indicated that a significant fraction of the variation in litter size, adjusted for female length, was due to consistent differences between individuals. Newborn weight varied within and among litters, but no relations between hatchling mass or mean egg mass in a litter and other traits were detected.
Size-adjusted reproductive performances remained constant during the course of this study, even though environmental conditions (weather factors, food availability) varied annually. Observed among year variations in reproductive characteristics were attributable to differences in the body size distributions of the adult females.