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.     

Geographic variation in reproduction and survival of kelp gulls Larus dominicanus vetula in southern Africa

Different populations of a species tend to vary in survival and reproduction, but the extent and scale of such spatial variation are poorly known. We estimated survival and clutch size of kelp gulls Larus dominicanus vetula across their entire African range. At this large geographic scale, we found no evidence for spatial variation in survival, and there was no variation in clutch size. However, there was considerable variation in clutch size among colonies within regions. Over the whole study, mean annual survival of juvenile and adult birds was 0.44 and 0.84, and mean clutch size was 2.2 eggs. A matrix population model showed that population growth was least sensitive to variation in clutch size, and the observed variation in clutch size could not fully account for the observed variation in population growth among colonies and regions. Our results thus suggest that dispersal and/or variation in survival (including egg/nestling survival) at a small spatial scale are also important for the spatial pattern of kelp gull population dynamics. These results are consistent with a metapopulation approach to spatial population dynamics.     

Is cell-mediated immunity related to the evolution of life-history strategies in birds?

According to life-history theory, the development of immune function should be balanced through evolutionary optimization of the allocation of resources to reproduction and through mechanisms that promote survival. We investigated interspecific variability in cell-mediated immune response (CMI), as measured by the phytohaemagglutinin (PHA) assay, in relation to clutch size, longevity and other life-history traits in 50 species of birds. CMI exhibited significant repeatability within species, and PHA responses in chicks were consistently stronger than in adults. Univariate tests showed a variety of significant relationships between the CMI of both chicks and adults with respect to size, development period and lifespan, but not clutch size or prevalence of blood parasites in adults. Multivariate analyses confirmed these patterns but independent variables were too highly correlated to isolate unique influences on CMI. The positive relationship of chick CMI to nestling period is further complicated by a parallel relationship of chick CMI to the age at testing. However, multivariate analysis showed that chick CMI varies uniquely with length of the nestling period. Adult CMI was associated with a strong life-history axis of body size, development rate and longevity. Therefore, adult CMI may be associated with prevention and repair mechanisms related to long lifespan, but it also may be allometrically related to body size through other pathways. Neither chick CMI nor adult CMI was related to clutch size, contradicting previous results linking parasite-related mortality to CMI and the evolution of clutch size (reproductive investment) in birds.     

Reproductive responses of birds to experimental food supplementation: a meta-analysis

Introduction

Food availability is an important environmental cue for animals for deciding how much to invest in reproduction, and it ultimately affects population size. The importance of food limitation has been extensively studied in terrestrial vertebrate populations, especially in birds, by experimentally manipulating food supply. However, the factors explaining variation in reproductive decisions in response to food supplementation remain unclear. By performing meta-analyses, we aim to quantify the extent to which supplementary feeding affects several reproductive parameters in birds, and identify the key factors (life-history traits, behavioural factors, environmental factors, and experimental design) that can induce variation in laying date, clutch size and breeding success (i.e., number of fledglings produced) in response to food supplementation.

Results

Food supplementation produced variable but mostly positive effects across reproductive parameters in a total of 201 experiments from 82 independent studies. The outcomes of the food effect were modulated by environmental factors, e.g., laying dates advanced more towards low latitudes, and food supplementation appeared not to produce any obvious effect on bird reproduction when the background level of food abundance in the environment was high. Moreover, the increase in clutch size following food addition was more pronounced in birds that cache food, as compared to birds that do not. Supplementation timing was identified as a major cause of variation in breeding success responses. We also document the absence of a detectable food effect on clutch size and breeding success when the target species had poor access to the feed due to competitive interactions with other animals.

Conclusions

Our findings indicate that, from the pool of bird species and environments reviewed, extra food is allocated to immediate reproduction in most cases. Our results also support the view that bird species have evolved different life-history strategies to cope with environmental variability in food supply. However, we encourage more research at low latitudes to gain knowledge on how resource allocation in birds changes along a latitudinal gradient. Our results also emphasize the importance of developing experimental designs that minimise competition for the supplemented food and the risk of reproductive bottle-necks due to inappropriate supplementation timings.

     

THE EVOLUTION OF MATERNAL INVESTMENT IN LIZARDS: AN EXPERIMENTAL AND COMPARATIVE ANALYSIS OF EGG SIZE AND ITS EFFECTS ON OFFSPRING PERFORMANCE

I used comparative and experimental analysis of egg size in a Sceloporus lizard to examine a fundamental tenet of life-history theory: the presumed trade-offs among offspring number, offspring size, and performance traits related to offspring size that are likely to influence fitness. I analyzed latitudinal and elevational patterns of egg life-history characteristics among populations and experimentally manipulated egg size and hatchling size by removing yolk from the eggs to examine the causal bases of population differences in offspring traits. Mean clutch size among populations increased to the north (seven vs. 12 eggs/clutch, California vs. Washington), whereas egg size decreased (0.65 g vs. 0.40 g). The elevational patterns in southern California paralleled the latitudinal trends. Several offspring life-history traits that are correlated with egg size also varied geographically; these traits included incubation time, hatchling size, growth rate, and hatchling sprint performance. Hatchling viability of experimentally reduced eggs was remarkably high (~70%), even when up to 50% of the yolk was removed. The experimentally reduced eggs and hatchlings demonstrated the degree to which size influences each of the offspring life-history traits considered. Northern eggs hatched sooner, in part because of their small size. Though growth rate is allometrically related to size within each population (i.e., smaller hatchlings grow faster on a mass-specific basis), population differences in growth rate, as measured in the laboratory, are likely to reflect genetic differentiation in the underlying physiology of growth. Moreover, smaller juveniles, because of experimental reduction, had slower sprint speeds than larger juveniles. The slower sprint speed of hatchlings from Washington compared to hatchlings from California is thus largely due to the fact that eggs are smaller in the Washington population. These results provide a basis for interpreting the evolutionary divergence of the suite of traits involved in the evolution of maternal investment per offspring in lizards. For example, evolutionary divergence in some offspring traits functionally related to size (e.g., sprint speed) may be constrained, relative to traits that are determined by other aspects of development or physiology (e.g., growth). I also discuss issues relating to the evolution of maternal investment that could be tested in laboratory and natural populations using experimentally reduced offspring.     

Life-history variation in closely related generalist predators living in the same habitat: a case study with three Cyclosa spiders

1. Generalist predators sharing similar food resources and phenologies as well as having no competitive interactions are expected to have a similar life-history pattern, but some closely related web spiders show different life-history traits. The present paper clarifies possible selection pressures affecting life-history traits of the three coexisting Cyclosa spiders and explores the significance of the life-history variation.
2. Cyclosa argenteoalba had lower daily survival rate and higher growth rate, C. sedeculata had higher daily survival rate and lower growth rate, and C. octotuberculata showed intermediate levels. This implies that the selection pressures these spiders experience differ appreciably even in the same habitat.
3. The significance of the life-history characteristics of the three species was evaluated by general life-history theories. Cyclosa argenteoalba showed distinguishing reproductive traits: shorter time to maturation, larger reproductive effort, larger relative clutch size, decreased clutch size with the number of clutches, and smaller egg size. These characteristics may have evolved in response to the larger ratio of juvenile to adult survivorship. Cyclosa octotuberculata had a clutch size much larger than the other two species, but relative clutch sizes accounting for body size were similar between C. octotuberculata and C. sedeculata . Also, the two species showed a similar time to maturation despite having different selection pressures. Probably, higher growth rate compensates for lower survivorship, leading to the similarity in some reproductive traits.     

Life-history traits of two Mediterranean lizard populations: a possible example of countergradient covariation

The trade-off between clutch and offspring size, which is a central topic in life-history research, is shaped by natural selection to maximize the number of surviving offspring, but it also depends on the resources available for reproduction. Conspecific populations living in different environments may differ in adult body size, clutch mass, clutch size, offspring size, and/or post-natal growth rates, due either to phenotypic plasticity or to local adaptation. Here, we compare these traits and their relationships between two populations of the lizard Psammodromus algirus separated by a 600-m altitudinal gradient. We used a common garden design to control incubation temperature and food availability, with two different feeding treatments. Females were larger at the high-elevation site. Although SVL-adjusted clutch mass did not differ between populations, high-elevation females laid more but smaller eggs than low-elevation ones. Hatchlings were larger at lower elevation. Our common garden experiment revealed that low-elevation hatchlings grew faster than high-elevation hatchlings under both feeding treatments. However, higher food availability at higher altitude allows high-elevation lizards to grow faster and attain larger adult sizes, especially in the case of females. The two key adaptations of low-elevation lizards, large eggs and hatchlings and the ability to grow rapidly after hatching, are likely to enhance survival in low-productivity Mediterranean lowlands. Our data support the hypothesis that the reproductive strategies of these populations provide an example of countergradient variation, because the genotypes that encode for fast growth and large body size occurred in low food availability habitats where juveniles grew slowly and attained small adult sizes.     

Covariation in Insular Life-history Traits of the Rice Frog(Fejervarya limnocharis)in Eastern China

Although shifts in life-history traits of insular vertebrates, as compared with mainland populations, have been observed in many taxa, few studies have examined the relationships among individual life-history traits on islands. Lifehistory theory also predicts that there is a trade-off between body size and reproductive effort, and between egg size and clutch size. We surveyed the rice frog, Fejervarya limnocharis, on 20 islands within the Zhoushan Archipelago and two nearby sites on the mainland of China to compare differences in life-history traits and to explore relationships among those traits. Rice frog females reached a greater body size on half of the smaller islands among the total 20 surveyed islands, and larger egg size, decreased clutch size and reduced reproductive effort on most of the islands when compared to the two mainland sites. Insular body size was negatively correlated with reproductive effort. There was a negative correlation between egg size and clutch size. Results suggest that life-history theory provides a good explanation for co-variation between body size and reproductive effort, and between egg size and clutch size in rice frogs on the islands.     

Intraspecific Variation of Reproductive Traits in a Mediterranean Lizard: Clutch, Population, and Lineage Effects

Widely distributed terrestrial ectotherms from the southern European peninsulas show patterns of subdivision (related to isolation in temperate refugia) that allow us to test the relative importance of phylogeographic lineage, population of origin and familial effects as sources of variation for life-history traits. We collected gravid females from 15 geographically separated populations of the lacertid lizard Psammodromus algirus, a widely distributed species with well differentiated eastern and western lineages. We incubated eggs under two treatments of constant (28°C) and fluctuating (28 ± 4°C) temperature, and we examined clutch, population, and lineage effects on several traits of females, eggs, and hatchlings. Incubation time was mainly explained by differences between lineages, but it was also influenced by population and female effects. Within each lineage, incubation was shorter at cooler and wetter sites, and for a given climate it was shorter for eastern than for western populations, suggesting that countergradient variation has evolved independently in the two lineages. Female size, clutch size, and relative fecundity were primarily influenced by inter-population differences, a pattern that seemed attributable to environmental differences in productivity, because mean female size was positively correlated with a gradient of increasing precipitation and decreasing temperature. Clutch size was mainly, but not entirely, dependent on female SVL, suggesting both a proximate effect of local conditions and intrinsic differences among populations. Females from drier and warmer sites produced larger hatchlings. Mean egg mass was mainly determined by familial effects. Eggs incubated at a constant temperature hatched earlier than did their siblings incubated at fluctuating temperatures, a fact that could be explained by considering that in Mediterranean environments developmental rate might increase at a lower speed above average incubation temperature than it does decrease below it.     

Adaptive inter-population differences in blue tit life-history traits on Corsica

Few studies of natural populations have investigated how phenotypic variation across populations relates to key factors in the environment and landscape structure. In the blue tits of southern France, inter-population differences in reproductive life-history traits (e.g. laying date and clutch size) are small, whatever the timing of maximum caterpillar availability, a key factor for offspring survival in tits. These small differences are attributed to gene flow between local populations occupying different habitat types. In contrast, in blue tits on the island of Corsica, we noted large differences in reproductive life-history traits between two populations, where each population is synchronized with the peak-date of caterpillar abundance. These occur over a short geographical distance (25km). Considering our study within a framework of long-term population studies in tits, our results support the hypothesis that different blue tit populations on Corsica show adaptive differences in life-history traits, and suggest that landscape structure at a small spatial scale can have profound effects on adaptive between-population differentiation in life-history traits that are closely linked with fitness.     

Interspecific associations between circulating antioxidant levels and life-history variation in birds

Antioxidants play an important role in protecting tissues against aging-associated oxidative damage and are thus prime candidates for relating physiological mechanisms to variation in life histories. We measured total antioxidant capacity, antioxidant response to stress, and levels of uric acid, vitamin E, and four carotenoids in 95 avian species, mostly passerines from Michigan or Panama. We compared antioxidant measures to seven variables related to life histories (clutch size, survival rate, incubation period, nestling period, basal metabolic rate, body mass, and whether the species lived in a tropical or temperate climate). Life-history-related traits varied over at least three statistically independent axes. Higher antioxidant levels were generally characteristic of more rapid development, lower survival rate, smaller body size, larger clutch size, and higher mass-adjusted metabolic rate, but the relationships of particular antioxidants with individual life-history traits showed considerable complexity. Antioxidant-life history associations differed between tropical and temperate species and varied with respect to taxonomic sampling. Vitamin E showed few relationships with life-history traits. Overall, our results partly support the hypothesis that antioxidant levels evolve to mirror free radical production. Clearly, however, the complex patterns of physiological diversification observed here result from the interplay of many factors, likely including not just investment in somatic maintenance but also phylogenetic constraint, diet, and other aspects of ecology.     

Nestling diet,secondary sexual traits and fitness in the zebra finch

We examined the effect of nestling diet quality on a suite of physiological, morphological and life-history traits in adult male zebra finches, Taeniopygia guttata. Compared with birds reared on a supplemented diet, nestlings reared on a seed-only diet showed a reduced rate of growth and reduced cell-mediated immune function as measured by an in vivo response to a T lymphocyte-dependent mitogen. There were no differences between birds reared on the two diets in any of the following adult traits: body size, primary sexual traits (testes mass, numbers of stored sperm, sperm function, velocity and morphology), secondary sexual traits (beak colour and song rate), serological traits or immunological traits. The only differences we detected were a lower body mass and a greater proportion of individuals with plumage abnormalities among those reared on a seed-only diet (this latter effect was transient). The fact that male zebra finches reared on a seed-only diet were, as adults, virtually indistinguishable from those reared on a supplemented diet, despite having reduced growth and immune function as nestlings, demonstrates that they subsequently compensated through the differential allocation of resources. Our results indicate that differential allocation is costly in terms of fitness since birds reared on a seed-only diet experienced a significantly greater mortality rate than those reared on a supplemented diet. This in turn suggests the existence of a trade-off between the development of traits important for reproduction, such as primary and secondary sexual traits and longevity.     

Fitness cost of incubation in great tits (Parus major) is related to clutch size

Life-history theory predicts that parents produce the number of offspring that maximizes their fitness. In birds, natural selection on parental decisions regarding clutch size may act during egg laying, incubation or nestling phase. To study the fitness consequences of clutch size during the incubation phase, we manipulated the clutch sizes during this phase only in three breeding seasons and measured the fitness consequences on the short and the long term. Clutch enlargement did not affect the offspring fitness of the manipulated first clutches, but fledging probability of the subsequent clutch in the same season was reduced. Parents incubating enlarged first clutches provided adequate care for the offspring of their first clutches during the nestling phase, but paid the price when caring for the offspring of their second clutch. Parents that incubated enlarged first clutches had lower local survival in the 2 years when the population had a relatively high production of second clutches, but not in the third year when there was a very low production of second clutches. During these 2 years, the costs of incubation were strong enough to change positive selection, as established by brood size manipulations in this study population, into stabilizing selection through the negative effect of incubation on parental fitness.     

A Review for Life-history Traits Variation in Frogs Especially for Anurans in China

Environmental variation can promote differentiation in life-history traits in species of anurans. Increased environmental stress usually results in larger age at sexual maturity, older mean age, longer longevity, slower growth, larger body size, and a shift in reproductive allocation from offspring quantity to quality, and a stronger trade-off between offspring size and number. However, previous studies have suggested that there are inconsistent geographical variations in life-history traits among anuran species in China. Hence, we here review the intraspecific patterns and differences in life-history traits(i.e., egg size, clutch size, testes size, sperm length, age at sexual maturity, longevity, body size and sexual size dimorphism) among different populations within species along geographical gradients for anurans in China in recent years. We also provide future directions for studying difference in sperm performance between longer and shorter sperm within a species through transplant experiments and the relationships between metabolic rate and brain size and life-history.     

Trade-offs in the life history and energy budget of the parthenogenetic collembolan Folsomia candida (Willem)

Three clones of Folsomia candida from different locations in Europe were compared in four experiments investigating genetic and phenotypic correlations between life-history traits. The first three experiments focused on the effects of food type, clone and temperature on traits associated with the first clutch. Differences in clutch size between clones and treatments were almost completely attributable to body size. Clones differed in length of the juvenile period, but the difference decreased at low temperatures. Age and weight at first reproduction were negatively correlated in the food type experiment and positively correlated in the temperature experiment, an often-encountered result for which no general explanation is as yet available. In the temperature experiment egg size variation was considerable, and was highest at low temperatures. The fourth experiment, with two clones at two feeding levels, aimed at finding trade-offs, in particular between reproduction and survival. It was hypothesized that higher fecundity led to increased scenescence through a higher metabolic rate. The trade-off was clearly present among the clones: one combined fast growth, late reproduction and high lifetime fecundity with lower survival, while in the other the relation between these traits was opposite. The proposed mechanism, however, was not confirmed, as no difference in metabolic rate was found. The effect of food level was too small to result in significant differences in the life-history traits in either of the clones.     

Clonal variation in the survival and reproduction of Daphniapulicaria under low-food stress

1. Survivorship and reproductive parameters of eight Daphnia pulicaria clones were evaluated in a life-table experiment under four dietary regimes ranging from food limitation to complete food deprivation.
2. D . pulicaria exhibited both diet-independent interclonal differences and genotype–environment interactions in life-span.
3. Reproduction ceased entirely or nearly so under the three treatments with lowest food availability. Under life-long food limitation, differences between clones were evident in clutch sizes and age at first reproduction. Some clones produced ephippia at a markedly higher frequency than others under food-limiting conditions.
4. For most of the life-history traits evaluated, within-lake variation among clones exceeded differences between the two lake populations they represented.
5. Intraspecific variations in response to periods of food deprivation and to extended quantitative or qualitative food-limitation could affect the clonal composition of Daphnia populations. They are therefore of potential importance in determining the effect of altered phytoplankton assemblages on zooplankton communities.     

Genetic and environmental components of growth in nestling blue tits (Parus caeruleus)

We investigated the effect of brood‐size mediated food availability on the genetic and environmental components of nestling growth in the blue tit (Parus caeruleus), using a cross‐fostering technique. We found genetic variation for body size at most nestling ages, and for duration of mass increase, but not of tarsus growth. Hence, nestling growth in our study population seems to have the potential to evolve further. Furthermore, significant genotype–environment interactions indicated heritable variation in reaction norms of growth rates and growth periods, i.e. that our study population had a heritable plasticity in the growth response to environmental conditions. The decreasing phenotypic variance with nestling age indicated compensatory growth in all body traits. Furthermore, the period of weight increase was longer for nestlings growing up in enlarged broods, while there was no difference to reduced broods in the period of tarsus growth. At fledging, birds in enlarged broods had shorter tarsi and lower weights than birds in reduced broods, but there was no difference in wing length or body condition between the two experimental groups. The observed flexibility in nestling growth suggests that growing nestlings are able to respond adaptively to food constraint by protecting the growth of ecologically important traits.     

Clonal variation in the survival and reproduction of Daphniapulicaria under low-food stress

1. Survivorship and reproductive parameters of eight Daphnia pulicaria clones were evaluated in a life-table experiment under four dietary regimes ranging from food limitation to complete food deprivation.
2. D . pulicaria exhibited both diet-independent interclonal differences and genotype–environment interactions in life-span.
3. Reproduction ceased entirely or nearly so under the three treatments with lowest food availability. Under life-long food limitation, differences between clones were evident in clutch sizes and age at first reproduction. Some clones produced ephippia at a markedly higher frequency than others under food-limiting conditions.
4. For most of the life-history traits evaluated, within-lake variation among clones exceeded differences between the two lake populations they represented.
5. Intraspecific variations in response to periods of food deprivation and to extended quantitative or qualitative food-limitation could affect the clonal composition of Daphnia populations. They are therefore of potential importance in determining the effect of altered phytoplankton assemblages on zooplankton communities.     

Are incubation costs in female pied flycatchers expressed in humoral immune responsiveness or breeding success?

Although clutch size variation has been a key target for studies of avian life history theory, most empirical work has only focused on the ability of parents to raise their altricial young. In this study, we test the hypothesis that costs incurred during incubation may be an additional factor constraining clutch size in altricial birds. In the pied flycatcher (Ficedula hypoleuca), we manipulated the incubation effort of the female by enlarging and reducing clutch sizes. To manipulate incubation effort only, the original clutch sizes were restored shortly after hatching. We found that fledging success was lower among broods whose clutches were enlarged during incubation. There was, however, no effect of manipulation on female body condition or on their ability to mount a humoral immune response to diphtheria or tetanus toxoid during the incubation or nestling provisioning period. Instead, we found that the original clutch size was related to the immune response so that females with seven eggs had significantly lower primary antibody responses against tetanus compared to those with six eggs. Our results suggest that incubating females are not willing to jeopardise their own condition and immune function, but instead pay the costs of incubating a larger clutch by lower offspring production. The results support the view that costs of producing and incubating eggs may be substantial and hence that these costs are likely to contribute to shaping the optimal clutch size in altricial birds.