Costs of reproduction in a tropical invariant-clutch producing lizard (Carlia rubrigularis)

Reproduction involves costs and benefits, whereby increased expenditure into current reproduction can reduce future reproductive success. Costs of reproduction, often assessed using the reduction in locomotor speed, have become well established in temperate-zone lizard species. However, substantial differences in biotic conditions and life-history traits between temperate- and tropical-zone lizards suggest that such costs may be different or of less importance for tropical species. This study examined the effects of reproduction on locomotor speed in the tropical invariant-clutch producing lizard Carlia rubrigularis . Counter to predictions and despite a low relative clutch mass (RCM), gravid and post-oviposition females experienced a reduction in locomotor speed with a physiological basis that was unrelated to the level of reproductive investment. In addition, gravid and postoviposition females exhibited locomotor speeds that were inversely related to the timing of oviposition and approached the speed of non-reproductive females after 3 weeks of oviposition. These results suggest that in addition to RCM, selection may act to reduce the period of recovery in species with extended reproductive seasons and which make numerous bouts of reproduction, such as C. rubrigularis .     

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.     

SEVERE COSTS OF REPRODUCTION PERSIST IN ANOLIS LIZARDS DESPITE THE EVOLUTION OF A SINGLE‐EGG CLUTCH

A central tenet of life‐history theory is that investment in reproduction compromises survival. We tested for costs of reproduction in wild brown anoles (Anolis sagrei) by eliminating reproductive investment via surgical ovariectomy and/or removal of oviductal eggs. Anoles are unusual among lizards in that females lay single‐egg clutches at frequent intervals throughout a lengthy reproductive season. This evolutionary reduction in clutch size is thought to decrease the physical burden of reproduction, but our results show that even a single egg significantly impairs stamina and sprint speed. Reproductive females also suffered a reduction in growth, suggesting that the cumulative energetic cost of successive clutches constrains the allocation of energy to other important functions. Finally, in each of two separate years, elimination of reproductive investment increased breeding‐season survival by 56%, overwinter survival by 96%, and interannual survival by 200% relative to reproductive controls. This extreme fitness cost of reproduction may reflect a combination of intrinsic (i.e., reduced allocation of energy to maintenance) and extrinsic (i.e., increased susceptibility to predators) sources of mortality. Our results provide clear experimental support for a central tenet of life‐history theory and show that costs of reproduction persist in anoles despite the evolution of a single‐egg clutch.     

Lifetime reproductive effort

In a 1966 American Naturalist article, G. C. Williams initiated the study of reproductive effort (RE) with the prediction that longer-lived organisms ought to expend less in reproduction per unit of time. We can multiply RE, often measured in fractions of adult body mass committed to reproduction per unit time, by the average adult life span to get lifetime reproductive effort (LRE). Williams's hypothesis (across species, RE decreases as life span increases) can then be refined to read "LRE will be approximately constant for similar organisms." Here we show that LRE is a key component of fitness in nongrowing populations, and thus its value is central to understanding life-history evolution. We then develop metabolic life-history theory to predict that LRE ought to be approximately 1.4 across organisms despite extreme differences in production and growth rates. We estimate LRE for mammals and lizards that differ in growth and production by five- to tenfold. The distributions are approximately normal with means of 1.43 and 1.41 for lizards and mammals, respectively (95% confidence intervals: 1.3-1.5 and 1.2-1.6). Ultimately, therefore, a female can only produce a mass of offspring approximately equal to 1.4 times her own body mass during the course of her life.     

Senescence and costs of reproduction in the life history of a small precocial species

Species following a fast life history are expected to express fitness costs mainly as increased mortality, while slow‐lived species should suffer fertility costs. Because observational studies have limited power to disentangle intrinsic and extrinsic factors influencing senescence, we manipulated reproductive effort experimentally in the cavy (Cavia aperea) which produces extremely precocial young. We created two experimental groups: One was allowed continuous reproduction (CR) and the other intermittent reproduction (IR) by removing males at regular intervals. We predicted that the CR females should senesce (and die) earlier and produce either fewer and/or smaller, slower growing offspring per litter than those of the IR group. CR females had 16% more litters during three years than IR females. CR females increased mass and body condition more steeply and both remained higher until the experiment ended. Female survival showed no group difference. Reproductive senescence in litter size, litter mass, and reproductive effort (litter mass/maternal mass) began after about 600 days and was slightly stronger in CR than IR females. Litter size, litter mass, and offspring survival declined with maternal age and were influenced by seasonality. IR females decreased reproductive effort less during cold seasons and only at higher age than CR females. Nevertheless, offspring winter mortality was higher in IR females. Our results show small costs of reproduction despite high reproductive effort, suggesting that under ad libitum food conditions costs depend largely on internal regulation of allocation decisions.     

The Edible Dormouse (<Emphasis Type="Italic">Glis glis</Emphasis>, Gliridae,Rodentia) in the Periphery of Its Distribution Range: Body Size and Life History Parameters

The body size, longevity, growth rate, and the intensity of breeding in a peripheral population of the edible dormouse from the Zhiguli Mountains are analyzed from the standpoint of life cycle theory. A breeding peculiarity of the species lies in periodically repeated years of reproductive failure, i.e., the total absence of young, seen throughout the species range. In reproductively successful years, anticipatory reproduction is observed, when the birth of posterity precedes a period of maximum food abundance. In the optimum of the distribution range, in Central and Western Europe, the mechanism of reproductive control in unfavorable years is based on the suppressing of the male reproductive activity at the beginning of the active season. In the Zhiguli population, natality regulation is rooted in mass resorption of embryos in the vast majority of females. In that population, the body size and weight are the minimum, while the difference from individuals from other populations studied may reach three times. The peculiarities of the edible dormouse biology in the peripheral population are related to the life cycle parameters. In contrast to the Central European populations, reproduction in the Zhiguli occurs every year, but in pregnant females, the death of embryos is observed in lean years. In males of that population, which are reproductively active every year, no significant costs of reproduction related to weight loss have been identified. The lower growth rate in comparison with more western European populations is shown in four age groups conventionally distinguishable based on morphometric parameters. In yearling males whose growth is not yet finished, the onset of the mating period depends on the body mass. Typically, the lifespan in the study population does not exceed four years, this being much shorter than in other populations where it lasts seven to nine years.     

Extended gestation with late‐autumn births in a cool‐climate viviparous gecko from southern New Zealand (Reptilia: Naultinus gemmeus)

Abstract Female viviparous lizards from temperate locations in the Southern Hemisphere (New Zealand, Tasmania (Australia), South Africa and South America) often have reproductive activity spanning many months of the year. In contrast, vitellogenesis and pregnancy are often confined to the spring/summer months in viviparous species from temperate zones of the Northern Hemisphere. An extreme Southern Hemisphere example is the nocturnal common gecko from New Zealand, Hoplodactylus maculatus (Gray 1845), in which females exhibit biennial reproduction with pregnancy lasting up to 14 months in a cool‐climate population. Here, we examined whether such an extended reproductive cycle also occurs in a diurnal species, the jewelled gecko Naultinus gemmeus (McCann 1955), at a similar latitude. Palpation was used to assess reproductive condition non‐invasively. In contrast to the nearby higher‐altitude population of H. maculatus, N. gemmeus reproduces annually. Vitellogenesis occurs from autumn to spring in both species, but pregnancy ends after about 7 months in N. gemmeus. Birth occurs in the seemingly unpropitious season of mid‐ to late autumn, a pattern that may be unique for lizards from cool‐temperate zones. We hypothesize that there are major differences between populations of N. gemmeus and H. maculatus with respect to survival of autumn‐born neonates and/or costs to females from remaining pregnant over winter. Museum specimens of N. gemmeus support anatomical inferences from palpation; they also suggest that vitellogenesis may begin before the end of pregnancy (which may be essential to completing each reproductive cycle within a year) and that some populations may show gestation in utero over winter, as in H. maculatus. Extended gestation appears to be a common response to cool climates for Southern Hemisphere lizards that have independently evolved viviparity.     

Effects of Limb Length,Body Mass,Gender, Gravidity,and Elevation on Escape Speed in the Lizard Psammodromus algirus

Most animals rely on their escape speed to flee from predators. Here, we test several hypotheses on the evolution of escape speed in the lizard Psammodromus algirus. We test that: (1) Longer limbs should improve speed sprint. (2) Heavier lizards should be impaired regarding their sprint speed ability, suggesting a trade-off between fat storage and escape capability. (3) Males should achieve faster speeds due to their higher exposure to predators. (4) Gravid females, with increased body mass, should perform lower speed than non-gravid females. And (5) there are inter-population differences in sprint speed across an elevational gradient. We measured lizards sprint speed in a lineal raceway in the laboratory, filming races in standardized conditions and then calculating their maximal speed. We found that hind limb length greatly determined maximal sprint speed, lizards with longer limbs being faster. In parallel, higher body masses reduced maximal speed, which points to a trade-off between fat storage and escaping capability. Sexual differences also arose, as males were faster than females, as a consequence of males having longer limbs. Regarding females, gravidity did not impair maximal sprint speed, suggesting adaptations which compensate for the increased body mass. Finally, we found no elevational trend in both limbs length and sprint speed. In any case, this study suggests that selection on escape capacity may cast morphological evolution, and affect other life-history traits, such as fat storage and reproduction.     

The evolution of locomotor morphology, performance, and anti-predator behaviour among populations of Leiocephalus lizards from the Dominican Republic

Most ecomorphological studies use a comparative approach to examine adaptation by studying variation among species. A question of considerable interest is whether ecomorphological patterns observed among species also exist at the population level. We studied variation in morphology, performance, and behaviour in four populations of Leiocephalus personatus and two populations of Leiocephalus barahonensis in the Dominican Republic. We combined these data with measurements of predation intensity and habitat structure to test for convergence at the population level. We predicted that predation intensity would be higher in open habitats and that lizards in these habitats would have traits conferring higher predator evasion capacity (increased wariness, faster sprint speeds, and longer limbs). Principal components analysis suggests that sites tend to differ with respect to the abundance and spacing of low-lying vegetation (i.e. percentage of shrub cover and distance to nearest vegetation), but we did not detect any striking differences among sites in tail-break frequencies or attacks on clay lizard models. Consistent with predictions we find that in open habitats, lizards tend to have longer limbs, faster sprint speeds (relative to body size), and longer approach distances. These patterns corroborate findings in other ground-dwelling lizard species and indicate that they have evolved at least twice among populations of Leiocephalus lizards. The results of this study also suggest that these traits have evolved rapidly despite recent or ongoing gene flow.  © 2008 The Linnean Society of London, Biological Journal of the Linnean Society , 2008, 93 , 445–456.     

Sex, Reproductive Status, and Cost of Tail Autotomy via Decreased Running Speed in Lizards

Autotomy, voluntary shedding of body parts to permit escape, is a theoretically interesting defense because escape benefit is offset by numerous costs, including impaired future escape ability. Reduced sprint speed is a major escape cost in some lizards. We predicted that tail loss causes decreased speed in males and previtellogenic females, but not vitellogenic females already slowed by mass gain. In the striped plateau lizard, Sceloporus virgatus , adults of both sexes are subject to autotomy, and females undergo large increases in body condition (mass/length) during vitellogenesis. Time required for running 1 m was similar in intact autotomized males and previtellogenic females, but increased by nearly half after autotomy. Vitellogenic females were slower than other lizards when intact, but their speed was unaffected by autotomy. Following autotomy, speeds of all groups were similar. Thus, speed costs of autotomy vary with sex and reproductive condition: decreased running speed is not a cost of autotomy in vitellogenic females or presumably gravid females. Costs of autotomy are more complex than previously known. Speed and other costs might interact in unforseen ways, making it difficult to predict whether strategies to compensate for diminished escape ability differ with reproductive condition in females.     

Physiological change in an insular lizard population confirms the reversed island syndrome

Unpredictable environmental conditions and highly fluctuating population densities are believed to have produced a ‘reversed island syndrome’ (RIS) in an insular population of the Wall lizard on Licosa Islet, Italy. Several of the physiological, behavioural, and life‐history changes that constitute the RIS could result from positive selection on increased activity of melanocortins. For example, increased levels of α‐melanocyte‐stimulating hormone (MSH) should lead to increased investment in reproduction and increased immunocompetence in the island population. We tested the crucial assumption of this idea that plasma levels of α‐MSH in Licosa Islet lizards are elevated compared to those of the mainland relatives. We also tested for differences in reproductive effort between populations, by measuring plasma levels of 5‐α‐dihydrotestosterone in males and clutch mass in females. In addition, we assessed ectoparasite load as an indicator for the lizards’ resistance to environmental stress. In agreement with the RIS, we found that insular lizards exhibit higher α‐MSH levels, allocate more energy to reproduction, and have a reduced ectoparasite load compared to the nearest mainland population. © 2012 The Linnean Society of London, Biological Journal of the Linnean Society, 2012, ●● , ●●–●●.     

Life‐history differences across latitude in common side‐blotched lizards (Uta stansburiana)

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Mass- and density-dependent reproductive success and reproductive costs in a capital breeder

For capital breeders, mass may affect reproductive potential. Reproductive expenditure may reduce future reproductive potential, particularly when resources are scarce. To test the hypothesis that reproductive success and the costs of reproduction vary according to mass and population density, we analyzed 25 yr of data on bighorn ewes (Ovis canadensis). The number of adult females was first limited by yearly removals, then allowed to triple. We found no survival costs of reproduction for ewes aged 4-7 yr. For ewes aged 8-14 yr, survival was density dependent for barren ewes but not for ewes that weaned lambs. Failure to lamb was rare and negatively correlated with fertility the following year. At low population density, lactation had a negative effect on mass gain but had a limited reproductive cost. At high density, heavy ewes had higher reproductive success than light ewes, and the reproductive cost and somatic costs of reproduction increased. The cost of reproduction was greater for light than for heavy ewes. Survival of weaned lambs to 1 yr was affected by population density but not by maternal mass or previous reproductive success. In large mammals, manipulations of reproductive effort are problematic, but long-term monitoring of individual mass and reproductive success under varying conditions of resource availability can provide insights into the evolution of life histories.     

Optimal life‐history schedule in a metapopulation with juvenile dispersal

Previous models have predicted that when mortality increases with age, older individuals should invest more of their resources in reproduction and produce less dispersive offspring, as both their future reproductive value and their prospect of competing with their own sib decline. Those models assumed stable population sizes. We here study for the first time the evolution of age‐specific reproductive effort and of age‐specific offspring dispersal rate in a metapopulation with extinction‐recolonization dynamics and juvenile dispersal. Our model explores the evolutionary consequences of disequilibrium in the age structure of individuals in local populations, generated by disturbances. Life‐history decisions are then shaped both by changes with age in individual performances, and by changes in ecological conditions, as young and old individuals do not live on average in the same environments. Lower juvenile dispersal favours the evolution of higher reproductive effort in young adults in a metapopulation with extinction‐recolonization compared with a well‐mixed population. Contrary to previous predictions for stable structured populations, we find that offspring dispersal should generally increase with maternal age. This is because young individuals, who are overrepresented in recently colonized populations, should allocate more to reproduction and less to dispersal as a strategy to exploit abundant recruitment opportunities in such populations.     

Climate‐dependent costs of reproduction: Survival and fecundity costs decline with length of the growing season and summer temperature

Costs of reproduction are expected to vary with environmental conditions thus influencing selection on life‐history traits. Yet, the effects of habitat conditions and climate on trade‐offs among fitness components remain poorly understood. For 2–5 years, we quantified costs of experimentally increased reproduction in two populations (coastal long‐season vs. inland short‐season) of two long‐lived orchids that differ in natural reproductive effort (RE; 30 vs. 75% fruit set). In both species, survival costs were found only at the short‐season site, whereas growth and fecundity costs were evident at both sites, and both survival and fecundity costs declined with increasing growing season length and/or summer temperature. The results suggest that the expression of costs of reproduction depend on the local climate, and that climate warming could result in selection favouring increased RE in both study species.     

EVOLUTIONARY PATTERNS OF THE THERMAL SENSITIVITY OF SPRINT SPEED IN ANOLIS LIZARDS

I present evidence that the thermal sensitivity of sprint speed of Anolis lizards has evolved to match the activity body temperatures (T_b) experienced by local populations in nature. Anolis lizards from a range of altitudes in Costa Rica have limited thermoregulatory abilities and consequently have field T_b that differ substantially in median and interquartile distance (a measure of variability). Experimentally determined maximal sprint temperatures (T_b at which lizards run fastest) were positively correlated with median field T_b, and performance breadths (ranges of T_b over which lizards run well) were correlated with the variability (interquartile distance) of field T_b in the species I examined. Such correlations would be expected if the thermal sensitivity of sprint speed and field T_b had evolved together to improve the sprint performance of lizards in nature. Integration of laboratory and field studies indicates that several species of Anolis regularly experience impaired sprint speeds in the field, despite apparent evolutionary modification of their thermal physiologies. However, this impairment would have been more severe if the thermal sensitivities of sprint speed had not evolved. Data from other groups of lizards indicate that the thermal sensitivity of sprint speed has not evolved to match T_b of local populations (Hertz et al., 1983; Crowley, 1985). These lizards experience less variable T_b and less impairment of sprint speeds in the field than do the anoles. Thus, selection for modification of the thermal sensitivity of sprint speed might have been stronger for anoles than for other groups of lizards.     

Habitat quality and among-population differentiation in reproductive effort and flowering phenology in the perennial herb <Emphasis Type="Italic">Primula farinosa</Emphasis>

In heterogeneous environments, selection on life-history traits and flowering time may vary considerably among populations because of differences in the extent to which mortality is related to age or size, and because of differences in the seasonal patterns of resource availability and intensity of biotic interactions. Spatial variation in optimal reproductive effort and flowering time may result in the evolution of genetic differences in life-history traits, but also in the evolution of adaptive phenotypic plasticity. The perennial herb Primula farinosa occurs at sites that differ widely in soil depth and therefore in water-holding capacity, vegetation cover, and frost-induced soil movement in winter. We used data from eight natural populations and a common-garden experiment to test the predictions that reproductive allocation is negatively correlated with soil depth while age at first reproduction and first flowering date among reproductive individuals are positively correlated with soil depth. In the common-garden experiment, maternal families collected in the field were grown from seed and monitored for 5 years. In the field, reproductive effort (number of flowers in relation to rosette area) varied among populations and was negatively related to soil depth. In the common-garden experiment, among-population differences in age at first reproduction, and reproductive effort were statistically significant, but relatively small and not correlated with soil depth at the site of origin. Flowering time varied considerably among populations, but was not related to soil depth at the site of origin. Taken together, the results suggest that among-population variation in reproductive effort observed in the field largely reflects phenotypic plasticity. They further suggest that among-population differentiation in flowering time cannot be attributed to variation in environmental factors correlated with soil depth.     

The Burden of Motherhood: The Effect of Reproductive Load on Female Lizard Locomotor,Foraging, and Social Behavior

The costs of reproduction, involving demands associated with both current and future reproductive efforts, may place a substantial burden on females. However, animals may minimize these costs by modifying their behavior across the reproductive cycle. We examined the effects of reproductive load on three types of behavior (locomotion, foraging, and social displays) in green anole lizards (Anolis carolinensis) by comparing egg, follicle, and oviduct mass and volume with field observational data. We found that female locomotor and social display behaviors decreased as reproductive load increased, suggesting behavioral modification in these traits, but we detected no relationship between foraging and reproductive load. We also examined these relationships across eight Anolis species using a phylogenetically informed analysis and found no associations between the evolution of reproductive load and any of the three behaviors. These results suggest that the evolution of increased reproductive load is not associated with the interspecific variation in behavior across the anoles and may result from varying life history traits or selective ecological pressures across species.     

绞股蓝雌雄种群觅源行为和繁殖对策比较

绞股蓝（Ｇｙｎｏｓｔｅｍｍａ ｐｅｎｔａｐｈｙｌｌｕｍ）雌雄异株,种群性比偏雄。作者利用比较生态学方法,从行为生态学角度探讨相同生境中绞股蓝雌雄种群的觅源行为和繁殖对策,得到如下初步结果和结论：（１）绞股蓝雄性种群的主枝生物量比显著大于雌性种群,这意味着雄性种群的营养繁殖投资显著增加,而两性种群在其它结构中的生物量分配无显著差异;（２）雌性种群的叶面积比和单位叶面积比雄性种群显著增加,这与两性种群     

Geographic variation in the effects of heat exposure on maximum sprint speed and Hsp70 expression in the western fence lizard Sceloporus occidentalis

We examined whether western fence lizards Sceloporus occidentalis occurring in thermally divergent environments display differential responses to high temperature in locomotor performance and heat-shock protein (Hsp) expression. We measured maximum sprint speed in S. occidentalis from four populations at paired latitudes and elevations before and after exposure to an experimental heat treatment and then quantified hind-limb muscle Hsp70 expression. Lizards collected from northern or high-elevation collection sites suffered a greater reduction in sprint speed after heat exposure than lizards collected from southern or low-elevation sites. In addition, lizards from northern collection sites also exhibited an increase in Hsp70 expression after heat exposure, whereas there was no effect of heat exposure on Hsp70 expression in lizards from southern collection sites. Across all groups, there was a negative relationship between Hsp70 expression and sprint speed after thermal stress. This result is significant because (a) it suggests that an increase in Hsp70 alone cannot compensate for the immediate negative effects of high-temperature exposure on sprint speed and (b) it demonstrates a novel correlation between an emergent property at the intersection of several physiological systems (locomotion) and a cellular response (Hsp70 expression). Ultimately, geographic variation in the effects of heat on sprint speed may translate into differential fitness and population viability during future increases in global air temperatures.