Selection on body size and sexual size dimorphism differs between host species in a seed-feeding beetle

Sexual size dimorphism varies substantially among populations and species but we have little understanding of the sources of selection generating this variation. We used path analysis to study how oviposition host affects selection on body size in a seed-feeding beetle (Stator limbatus) in which males contribute large ejaculates (nuptial gifts) to females. Females use nutrients in these ejaculates for egg production. Male body size, which affects ejaculate size, affects female fecundity and is thus under fecundity selection similar in magnitude to the fecundity selection on female body size. We show that when eggs are laid on a host on which larval mortality is low (seeds of Acacia greggii) fecundity predicts fitness very well and fecundity selection is the major source of selection on both male and female adult size. In contrast, when eggs are laid on a host on which larval mortality is high (seeds of Parkinsonia florida) fecundity poorly predicts fitness such that fecundity selection is relaxed on both male and female size. However, because egg size affects larval mortality on this poor host (P. florida) there is selection on female size via the female size --> egg size --> fitness path; this selection via egg size offsets the reduction in fecundity selection on female, but not male, body size. Thus, differences in host suitability (due to differences in larval mortality) affect the relative importance of two sources of selection on adult body size; fecundity selection on both male and female body size is lower on the poor quality host (P. florida) relative to the high quality host (A. greggii) whereas selection on female body size via effects of egg size on offspring survival (body size --> egg size --> fitness) is greater on the poor quality host relative to the high quality host. Because selection via the egg size path affects only females the difference in larval survival between hosts shifts the relative magnitude of selection on female vs. male size. Researchers working on other study systems should be alerted to the possible importance of subtle, but consequential, indirect selection on their study organisms.     

Expression of a life-history trade-off in a seed beetle depends on environmental context

Within populations of the seed beetle Callosobruchus maculatus (F.) (Coleoptera: Bruchidae), female longevity is positively correlated with fecundity, both phenotypically and genetically. Yet manipulations of egg-laying rate (through deprivation of seeds or mates) consistently suggest a trade-off between these traits. In this study, females were exposed to four levels of seed availability and the relationship between fecundity and longevity was examined both across and within environments. The expected trade-off was observed across environments, as females with few or no seeds laid fewer eggs and lived longer than females with many seeds. Within environments, however, the relationship was inconsistent; individual longevity and fecundity were positively correlated when seeds were abundant, uncorrelated when seeds were few and negatively correlated when seeds were absent. Body mass at adult emergence was positively correlated with fecundity, but its effect was stronger when seeds were present than when they were absent. After the effects of mass were statistically controlled, longevity and fecundity remained positively correlated among females with many seeds and negatively correlated among those without seeds. Thus, even a single technique (phenotypic correlation) can produce divergent conclusions about the ‘cost of reproduction’ in insects. The reversal in the correlation appears to depend on variation in reproductive allocation; when seeds are scarce or absent, differences in reproductive effort among individuals may be large enough to overwhelm differences in resource acquisition.     

Genetic variation in male effects on female reproduction and the genetic covariance between the sexes

Abstract.— Males of many insect species increase the fecundity and/or egg size of their mates through the amount or composition of their nuptial gifts or ejaculate. The genetic bases of such male effects on fecundity or egg size are generally unknown, and thus their ability to evolve remains speculative. Likewise, the genetic relationship between male and female investment into reproduction in dioecious species, which is expected to be positive if effects on fecundity are controlled by at least some of the same genes in males and females, is also unknown. Males of the seed beetle Stator limbatus contribute large ejaculates to females during mating, and the amount of donated ejaculate is positively correlated with male body mass. Females mated to large males lay more eggs in their lifetime than females mated to small males. We describe an experiment in which we quantify genetic variation in the number of eggs sired by males (mated to a single female) and found that a significant proportion of the phenotypic variance in the number of eggs sired by males was explained by their genotype. Additionally, the number of eggs sired by a male was highly positively genetically correlated with his body mass. The between-sex genetic correlation, that is, the genetic correlation between the number of eggs sired by males and the number of eggs laid by females, was highly positive when eggs were laid on Acacia greggii seeds. This indicates that males that sire many eggs have sisters that lay many eggs. Thus, some of the genes that control male ejaculate size (or some other fecundity-enhancing factor) when expressed in males appear to control fecundity when expressed in females. We found no significant interaction between male and female genotype on fecundity.     

Egg Size Versus Number of Offspring Trade-Off: Female Age Rather Than Size Matters in a Domesticated Arctic Charr Population

Understanding how organisms adjust reproductive allocation trade-offs between offspring size versus number (OSN trade-off) is a central question in evolutionary biology. In organisms with indeterminate growth, changes in OSN according to maternal size or age have been reported in numerous taxa. The relative contribution of age and size remains largely unclear, as they are often highly correlated. In this study, we investigated how females adjust the offspring size versus number trade-off and analyzed the relative contribution of female age and size in a domesticated population of Arctic charr Salvelinus alpinus (Linnaeus, 1758) that exhibit large variation in size within five age classes. Our results show that the reproductive output (i.e. as measured by the clutch mass), was strongly correlated to female mass and age suggesting that the proportion of resources allocated to reproduction do not vary along lifetime. Egg mass and fecundity (egg number) increased with female mass overall. However, within an age class, larger females had higher fecundity but egg mass was poorly related to female mass. At the population level, a positive relationship was observed between fecundity and egg mass but within each class age the relation was negative revealing a OSN trade-off. Overall, our results show that, in our model Arctic charr population, allocation trade-off to reproduction and the way females allocate to egg mass and fecundity is largely determined by their age rather than mass.     

Do mothers producing large offspring have to sacrifice fecundity?

We artificially selected on egg size in a butterfly to study the consequences for fecundity, reproductive effort and offspring fitness. Correlated responses in either pupal mass, larval or pupal development time were virtually absent. Offspring size was positively related to fitness, but only partly traded off against fecundity. Rather, total reproductive effort (measured as fresh mass), egg water content and the decline of egg size with female age increased in the large-egg selected lines compared to either small-egg or control lines. Accounting for these effects showed that reproductive investment (in dry mass) was in fact similar across lines. Such mechanisms may enable increased investment in (early) offspring without a reduction in their number, revealing a much more complex picture than a simple trade-off between offspring size and number. Substantial variation among replicates suggests that there are different underlying mechanisms for change, rather than any single, unitary pathway.     

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.     

Artificial selection on male longevity influences age-dependent reproductive effort in the black field cricket Teleogryllus commodus

Although the trade-off between reproductive effort and longevity is central to both sexual selection and evolutionary theories of aging, there has been little synthesis between these fields. Here, we selected directly on adult longevity of male field crickets Teleogryllus commodus and measured the correlated responses of age-dependent male reproductive effort, female lifetime fecundity, and several other life-history traits. Male longevity responded significantly to five generations of divergent selection. Males from downward-selected lines commenced calling sooner and reached their peak calling effort at a younger age. They called more per night and, despite living less than half as long, called more overall than males selected for increased longevity. Females from the downward-selected lines lived significantly shorter lives than females from the upward-selected lines but still produced the same number of offspring. Nymph survival, development time, and body size and weight at eclosion did not show significant correlated response to selection on male longevity, despite evidence for substantial genetic variation in each of these traits. Collectively, our findings directly support the antagonistic pleiotropy model of aging and suggest an important role for sexual selection in the aging process.     

Life-history phenotypes in populations of Brachyrhaphis episcopi (Poeciliidae) with different predator communities

Variation among populations in extrinsic mortality schedules selects for different patterns of investment in key life-history traits. We compared life-history phenotypes among 12 populations of the live-bearing fish Brachyrhaphis episcopi. Five populations co-occurred with predatory fish large enough to prey upon adults, while the other seven populations lacked these predators. At sites with large predatory fish, both sexes reached maturity at a smaller size. Females of small to average length that co-occurred with predators had higher fecundity and greater reproductive allotment than those from populations that lacked predators, but the fecundity and reproductive allotment of females one standard deviation larger than mean body length did not differ among sites. In populations with large predatory fish, offspring mass was significantly reduced. In each population, fecundity, offspring size and reproductive allotment increased with female body size. When controlling for maternal size, offspring mass and number were significantly negatively correlated, indicating a phenotypic trade-off. This trade-off was non-linear, however, because reproductive allotment still increased with brood size after controlling for maternal size. Similar differences in life-history phenotypes among populations with and without large aquatic predators have been reported for Brachyrhaphis rhabdophora in Costa Rica and Poecilia reticulata (a guppy) in Trinidad. This may represent a convergent adaptation in life-history strategies attributable to predator-mediated effects or environmental correlates of predator presence.     

Female freshwater crayfish adjust egg and clutch size in relation to multiple male traits

Females may invest more in reproduction if they acquire mates of high phenotypic quality, because offspring sired by preferred partners may be fitter than offspring sired by non-preferred ones. In this study, we tested the differential maternal allocation hypothesis in the freshwater crayfish, Austropotamobius italicus, by means of a pairing experiment aimed at evaluating the effects of specific male traits (body size, chelae size and chelae asymmetry) on female primary reproductive effort. Our results showed that females laid larger but fewer eggs for relatively small-sized, large-clawed males, and smaller but more numerous eggs for relatively large-sized, small-clawed males. Chelae asymmetry had no effects on female reproductive investment. While the ultimate consequences of this pattern of female allocation remain unclear, females were nevertheless able to adjust their primary reproductive effort in relation to mate characteristics in a species where inter-male competition and sexual coercion may mask or obscure their sexual preferences. In addition, our results suggest that female allocation may differentially affect male characters, thus promoting a trade-off between the expression of different male traits.     

Factors affecting variation in the reproductive investment of female treefrogs,Hyla intermedia

In anurans, fecundity (clutch size) is the most important determinant of female reproductive success. We investigated three possible causes responsible for fecundity variation in female Italian treefrogs, Hyla intermedia, during four breeding seasons: (i) variation in morphological (body size and condition) and life-history (age) traits; (ii) variation in the tradeoff between the number and the size of eggs; (iii) seasonal effects and within-season differences in the timing of deposition. At the population level, we found no evidence for a tradeoff between the number and the size of eggs, because they both correlated positively with females’ body size. Conversely, neither age nor post-spawning body condition showed any effect on female reproductive investment. Independent of body size, we found no evidence for variation in reproductive effort among different breeding seasons, but strong evidence for a decrease of clutch size and an increase of egg size with the advancing of a breeding season. To test for the functional significance of the observed temporal variation in allocation strategy, we carried out a rearing experiment in semi-natural conditions on a random sample of ten clutches. The experiment showed a negative effect of clutch size and a positive effect of egg size on both tadpole growth and developmental rates, suggesting that reproductive investment, although constrained by body size, can be adjusted by females to the time of deposition to increase the chances of offspring survival.     

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.     

Hormonal manipulation of offspring number: maternal effort and reproductive costs

We used exogenous gonadotropin hormones to physiologically enlarge litter size in the bank vole (Clethrionomys glareolus). This method allowed the study design to include possible production costs of reproduction and a trade-off between offspring number and body size at birth. Furthermore, progeny rearing and survival and postpartum survival of the females took place in outdoor enclosures to capture salient naturalistic effects that might be present during the fall and early winter. The aim of the study was to assess the effects of the manipulation on the growth and survival of the offspring and on the reproductive effort, survival, and future fecundity of the mothers. Mean offspring body size was smaller in enlarged litters compared to control litters at weaning, but the differences disappeared by the winter. Differences in litter sizes disappeared before weaning age due to higher mortality in enlarged litters. In addition to the effects of the litter size, offspring performance was probably also influenced by the ability of the mother to support the litter. Experimental females had higher reproductive effort at birth, and they also tended to have higher mortality during nursing. Combined effects of high reproductive effort at birth and high investment in nursing the litter entailed costs for the experimental females in terms of decreased probability of producing a second litter and a decreased body mass gain. Thus, enlarged litter size had both survival and fecundity costs for the mothers. Our results suggest that the evolution of litter size and reproductive effort is determined by reproductive costs for the mothers as well as by a trade-off between offspring number and quality.     

Cascading costs of reproduction in female house wrens induced to lay larger clutches

In many species, females produce fewer offspring than they are capable of rearing, possibly because increases in current reproductive effort come at the expense of a female's own survival and future reproduction. To test this, we induced female house wrens (Troglodytes aedon) to lay more eggs than they normally would and assessed the potential costs of increasing cumulative investment in the three main components of the avian breeding cycle – egg laying, incubation and nestling provisioning. Females with increased clutch sizes reared more offspring in the first brood than controls, but fledged a lower proportion of nestlings. Moreover, nestlings of experimental females were lighter than those of control females as brood size and prefledging mass were negatively correlated. In second broods of the season, when females were not manipulated, experimental females laid the same number of eggs as controls, but experienced an intraseasonal cost through reduced hatchling survival and a lower number of young fledged. Offspring of control and experimental females were equally likely to recruit to the breeding population, although control females produced more recruits per egg laid. The reproductive success of recruits from broods of experimental and control females did not differ. The manipulation also induced interseasonal costs to future reproduction, as experimental females had lower fecundity than controls when breeding at least 2 years after having their reproductive effort experimentally increased. Finally, females producing the modal clutch size of seven eggs in their first broods had the highest lifetime number of fledglings.     

Survival costs of reproduction predict age-dependent variation in maternal investment

Life-history theory predicts that older females will increase reproductive effort through increased fecundity. Unless offspring survival is density dependent or female size constrains offspring size, theory does not predict variation in offspring size. However, empirical data suggest that females of differing age or condition produce offspring of different sizes. We used a dynamic state-variable model to determine when variable offspring sizes can be explained by an interaction between female age, female state and survival costs of reproduction. We found that when costs depend on fecundity, young females with surplus state increase offspring size and reduce number to minimize fitness penalties. When costs depend on total reproductive effort, only older females increase offspring size. Young females produce small offspring, because decreasing offspring size is less expensive than number, as fitness from offspring investment is nonlinear. Finally, allocation patterns are relatively stable when older females are better at acquiring food and are therefore in better condition. Our approach revealed an interaction between female state, age and survival costs, providing a novel explanation for observed variation in reproductive traits.     

Geographical variation in reproductive traits and trade‐offs between size and number of eggs in the king ratsnake,Elaphe carinata

We collected gravid king ratsnakes (Elaphe carinata) from three geographically separated populations in Chenzhou (CZ), Lishui (LS) and Dinghai (DH) of China to study the geographical variation in female reproductive traits and trade‐offs between the size and number of eggs. Not all reproductive traits varied among the three populations. Of the traits examined, five (egg‐laying date, post‐oviposition body mass, clutch size, egg mass and egg width) differed among the three populations. The egg‐laying date, ranging from late June to early August, varied among populations in a geographically continuous trend, with females at the most northern latitude (DH) laying eggs latest, and females at the most southern latitude (CZ) laying eggs earliest. Such a trend was less evident or even absent in the other traits that differed among the three populations. CZ and DH females, although separated by a distance of approximately 1100 km as the crow flies, were similar to each other in most traits examined. LS females were distinguished from CZ and DH females by the fact that they laid a greater number of eggs, but these were smaller. The egg size–number trade‐off was evident in each of the three populations and, at a given level of relative fecundity, egg mass was significantly greater in the DH population than in the LS population. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 104 , 701–709.     

Female reproductive characteristics of the Balkan wall lizard (<Emphasis Type="Italic">Podarcis taurica</Emphasis>) in the northwestern periphery of its range

We present data on the female reproductive traits of the Balkan wall lizard in the Deliblato Sand, a large continental sandland in the Pannonian area in the northwestern periphery of the species range. The clutch and egg characteristics of the population were investigated on the basis of clutches laid in laboratory conditions by gravid females captured in one locality. Balkan wall lizards produced at least two clutches in a breeding season. Individual females laid clutches of commonly two (range 1–4) eggs. The female body size had no effect on clutch and egg size. There was no trade-off between egg size and clutch size.     

密点麻蜥的两性异形和雌性繁殖

蜥蜴繁殖成功率与其形态特征有密切的关系。作者在内蒙古乌拉特后旗采集密点麻蜥(Eremias multio-cellata) ,定量研究该种形态特征的两性异形和雌体繁殖特征,检验与成体形态特征相关的两性繁殖成功率差异是否能促进两性异形的进化。密点麻蜥成体个体大小无显著的两性差异,但头部大小两性差异显著;雄性个体的头长和头宽均大于体长相同的雌性成体。繁殖雌体于五、六月份排卵;在实验室条件下,雌体在六月下旬至七月下旬之间产仔。该种雌体年产单窝仔,每窝2 -4仔。窝仔重与雌体体长呈正相关,但雌体体长仅能解释很少一部分(约19 %)窝仔重的变异。窝仔数和幼仔重均与雌体体长无关。幼仔重与相对生育力(相对于雌体体长的窝仔数)呈显著的负相关,表明该种蜥蜴存在后代数量-大小之间的权衡。密点麻蜥雄体和雌体向较大体型方向进化的选择压力均相对较弱,与成体头部大小相关的两性繁殖成功率的差异是导致该种蜥蜴头部大小两性异形进化的主要原因[动物学报52 (2) : 250 -255 , 2006]。     

Breeding phenology,variation in reproductive effort and offspring size in a tropical population of the woodlouse Porcellionides pruinosus

Summary Most species of woodlice in temperate habitats have discrete breeding seasons. It is hypothesised that breeding synchronises with favourable environmental conditions to maximise offspring growth and survivorship (Willows 1984). We measured the breeding phenology of a species introduced to a tropical environment, primarily to consider the assumption that life histories in the tropics will differ fundamentally from those in temperate habitats. In addition to breeding phenology we considered variation in reproductive effort between individual females and the division of this effort between the size and number of young.A continuous breeding phenology was observed in a synanthropic population of Porcellionides pruinosus within the tropics. Reproductive effort varied between months, showed a weak relationship with female size and was independent of female fecundity. Female sizefecundity relationships varied between samples and when the proportion of reproductive females was high size-fecundity slopes were steeper than at other times. Mean offspring size varied between months and there was a wide range in offspring size within broods. Offspring size was not related to female body mass, reproductive effort or fecundity; consequently brood mass increased linearly with an increase in fecundity. Increased reproductive effort goes into more rather than larger offspring.We propose that the continuous breeding in this population was the result of the constant presence of an environmental cue to reproduction evolved in temperate habitats. Continuous breeding is not necessarily equivocal to high individual reproductive success even though overall population growth may be rapid. However, variation in reproductive effort suggests that individuals respond to current environmental conditions on short time scales.     

Reproductive plasticity, ovarian dynamics and maternal effects in response to temperature and flight in Pararge aegeria

In nature, ovipositing females may be subjected to multiple extrinsic and intrinsic environmental factors simultaneously. To adequately assess a species response to environmental conditions during oviposition it may therefore be necessary to consider the interaction between multiple intrinsic and extrinsic factors simultaneously. Using the butterfly, Pararge aegeria, this study examined the combined effects of extrinsic (temperature and flight) and intrinsic (body mass and age) factors on ovarian dynamics, egg provisioning and reproductive output, and explored how these effects subsequently influenced offspring fitness when egg-stage development occurred in a low humidity environment. Both temperature- and flight-mediated plasticity in female reproductive output was observed, and there were strong temperature by flight interaction effects for the traits oocyte size and egg mass. As females aged, mean daily fecundity differed across temperature treatments, but not across flight treatments. Overall, temperature had more pronounced effects on ovarian dynamics than flight. Flight mainly influenced egg mass via changes in relative water content. A mismatch between the physiological response of females to high temperature and the requirements of their offspring had a negative impact on offspring fitness via effects on egg hatching success.     

Male genotype influences female reproductive investment in Drosophila melanogaster

In many species, males can influence the amount of resources their mates invest in reproduction. Two favoured hypotheses for this observation are that females assess male quality during courtship or copulation and alter their investment in offspring accordingly, or that males manipulate females to invest heavily in offspring produced soon after mating. Here, we examined whether there is genetic variation for males to influence female short-term reproductive investment in Drosophila melanogaster, a species with strong sexual selection and substantial sexual conflict. We measured the fecundity and egg size of females mated to males from multiple isofemale lines collected from populations around the globe. Although these traits were not strongly influenced by the male's population of origin, we found that 22 per cent of the variation in female short-term reproductive investment was attributable to the genotype of her mate. This is the first direct evidence that male D. melanogaster vary genetically in their proximate influence on female fecundity, egg size and overall reproductive investment.