Sex-related differences in reaction norms in the butterfly Lycaena tityrus (Lepidoptera: Lycaenidae)

We investigate sexual differences in reaction norms in directly developing individuals of the copper butterfly Lycaena tityrus predicted from sexual selection theory. As recent studies on butterflies revealed a high degree of adaptive plasticity in growth and development, which may undermine the basic trade‐offs assumed in life‐history theory, we focus on effects of temperature, trying to drive growth rates to their physiological upper limit and thus disclosing otherwise potentially concealed responses. Development time strongly depended on temperature, leading in accordance with a central assumption in life‐history theory to a larger size at low temperatures, and vice versa. At all temperatures larval development time of males was significantly shorter compared to females, as was predicted by protandry theory. This was partially due to an invariably higher growth rate of males. However, sexes responded in different ways to developmental time constraints caused by increasing temperatures. Despite the shorter larval time of males, both sexes achieved similar body sizes at lower temperatures, because males avoided a reduction in weight due to plastic growth. At high temperatures, in contrast, males were forced to make a trade‐off in which they favoured early emergence over large size, leading to a dramatic weight loss. Weight of females, however, remained similar throughout showing no trade‐off. These different reaction norms reflect divergent selective pressures acting on males and females, which can be explained in relation to the reproductive system. The strong selection for early emergence in males is likely to be due to monandry, discrete non‐overlapping generations (as was already predicted by theory), and territoriality, because prior ownership of a territory seems to be a major advantage for successful reproduction. On the other hand, the preference of females for large body size was expected due to the close relationship between this trait and fecundity. Thus, our results highlight the extraordinary importance of the specific reproductive system, which can influence central life‐history traits in manifold ways.     

Sexual differences in larval life history traits of acanthocephalan cystacanths

Sexual differences in life history traits, such as size dimorphism, presumably arise via sexual selection and are most readily observed in adults. For complex life-cycle parasites, however, sexual selection may also have consequences for larval traits, e.g., growth in intermediate hosts. Two acanthocephalan species (Acanthocephalus lucii and Echinorhynchus borealis) were studied to determine, whether larval life histories differ between males and females. The size of female A. lucii cystacanths had a much stronger relationship with intermediate host size than males, suggesting females invest more in growth and are consequently more limited by resources. No relationship between host size and cystacanth size was observed for E. borealis. For both species, female cystacanths survived longer in a culture medium composed entirely of salts, which could suggest that females have greater energy reserves than males. A comparative analysis across acanthocephalan species indicated that sexual size dimorphism at the adult stage correlates with cystacanth dimorphism. However, the relationship was not isometric; cystacanths do not reach the same level of sexual dimorphism as adults, possibly due to resource constraints. Our results suggest that larval life histories diverge between males and females in some acanthocephalans, and this is seemingly a consequence of sexual selection acting on adults.     

Correlated responses to artificial body size selection in growth, development, phenotypic plasticity and juvenile viability in yellow dung flies

Most life history traits are positively influenced by body size, whereas disadvantages of large body size are poorly documented. To investigate presumed intrinsic costs of large size in the yellow dung fly (Scathophaga stercoraria; Diptera: Scathophagidae), we established two replicates each of three body size laboratory selection lines (small, control and large; selection on males only), and subjected flies of the resulting extended body size range to various abiotic stresses. Response to selection was symmetrical in the small and large lines (realized h(2) = 0.16-0.18). After 24 generations of selection body size had changed by roughly 10%. Female size showed a correlated response to selection on male size, whereas sexual size dimorphism did not change. Development time also showed a correlated response as, similar to food limited flies, small line flies emerged earlier at smaller body size. At the lowest larval food limit possible, flies of all lines emerged at the same small body size after roughly the same development time; so overall phenotypic plasticity in body size and development time strongly increased following selection. Juvenile mortality increased markedly when food was extremely limited, large line flies showing highest mortality. Winter frost disproportionately killed large (line) flies because of their longer development times. Mortality at high temperatures was high but size-selective effects were inconsistent. In all environments the larger males suffered more. Initial growth rate was higher for males and at unlimited food. Small line individuals of both sexes grew slowest at unlimited larval food but fastest at limited larval food, suggesting a physiological cost of fast growth. Overall, extension of the natural body size range by artificial selection revealed some otherwise cryptic intrinsic juvenile viability costs of large size, mediated by longer development or faster growth, but only in stressful environments.     

Resource availability modulates the effect of body size on reproductive development

Within-species variation in animal body size predicts major differences in life history, for example, in reproductive development, fecundity, and even longevity. Purely from an energetic perspective, large size could entail larger energy reserves, fuelling different life functions, such as reproduction and survival (the “energy reserve” hypothesis). Conversely, larger body size could demand more energy for maintenance, and larger individuals might do worse in reproduction and survival under resource shortage (the “energy demand” hypothesis). Disentangling these alternative hypotheses is difficult because large size often correlates with better resource availability during growth, which could mask direct effects of body size on fitness traits. Here, we used experimental body size manipulation in the freshwater cnidarian Hydra oligactis, coupled with manipulation of resource (food) availability to separate direct effects of body size from resource availability on fitness traits (sexual development time, fecundity, and survival). We found significant interaction between body size and food availability in sexual development time in both males and females, such that large individuals responded less strongly to variation in resource availability. These results are consistent with an energy reserve effect of large size in Hydra. Surprisingly, the response was different in males and females: small and starved females delayed their reproduction, while small and starved males developed reproductive organs faster. In case of fecundity and survival, both size and food availability had significant effects, but we detected no interaction between them. Our observations suggest that in Hydra, small individuals are sensitive to fluctuations in resource availability, but these small individuals are able to adjust their reproductive development to maintain fitness.     

Two sexes respond equally to food restriction in a sexually dimorphic but not body mass dimorphic jumping spider

Natural selection favors animals that evolve developmental and behavioral responses that buffer the negative effects of food restrictions. These buffering responses vary both between species and within species. Many studies have shown sex‐specific responses to environmental changes, usually in species with sexual size dimorphism (SSD), less found in species with weak or no SSD, which suggests that sizes of different sexes are experiencing different selections. However, previous studies usually investigated development and behavior separately, and the balanced situation where males and females of sexually dimorphic species respond in the same way to food restriction remains little known. Here, we investigated this in Phintelloides versicolor (Salticidae) that presents sexual dimorphism in color and shape but weak SSD. We examined whether food restriction induced the same responses in males and females in development duration, adult body size and weight, daily time allocated to foraging, and hunting. We found food restriction induced similar responses in both sexes: both exhibited longer development duration, smaller adult body size and weight, higher probability of staying outside nests and noticing prey immediately, and higher hunting success. However, there were sexual differences regardless of food condition: females showed faster development, smaller adult body size, higher probability of staying outside of nests, and higher hunting success. These indicated the differential selection on male and female sizes of P. versicolor could be under a balanced situation, where males and females show equal developmental and behavioral plasticity to environmental constraints.     

Sexual size dimorphism in the ontogeny of the solitary predatory wasp Symmorphus allobrogus (Hymenoptera: Vespidae)

Sex-specific patterns of individual growth, resulting in sexual size dimorphism (SSD), are a little studied aspect of the ontogeny related to the evolutionary history and affected by the ecology of a species. We used empirical data on the development of the predatory wasp Symmorphus allobrogus (Hymenoptera, Vespidae) to test the hypotheses that sexual differences of growth resulting in the female-biased SSD embrace the difference in (1) the egg size and the starting size of larva, (2) the larval development duration, and (3) the larval growth rate. We found that eggs developing into males and females have significant differences in size. There was no significant difference between the sexes in the duration of larval development. The relative growth rate and the food assimilation efficiency of female larvae were significantly higher than compared to those of male larvae. Thus, the SSD of S. allobrogus is mediated mainly by sexual differences in egg size and larval growth rate.     

Adaptive sex-specific life history plasticity to temperature and photoperiod in a damselfly

We investigated four predictions about how temperature, photoperiod and sex affect the life history plasticity and foraging activity of a damselfly. (i) As predicted, increased temperatures increased foraging activity and growth rates, but in contrast with the prediction, late photoperiod (high time stress) did not affect foraging activity and growth rate. (ii) Unexpectedly, the increase in growth rate at increasing temperatures was not larger under high time stress. (iii) As predicted, age and size at emergence decreased at higher temperatures and at the late photoperiod. Temperature-induced life history shifts were direct or the result of behavioural growth mediation depending on the temperature range. Photoperiod-induced life history shifts were direct. (iv) As predicted, males emerged before females but at a smaller size. The degree of sexual size dimorphism was influenced by the joint effects of temperature and photoperiod. We could only detect genetic variation in size plasticity to photoperiod. The match between the sex-specific life history responses to temperature and photoperiod and predictions by relevant optimality models suggests adaptive life history plasticity to these variables.     

Patterns in Size,Sex Ratio and Time at Emergence in a South Swedish Population of Sympetrum sanguineum (Odonata)

Differences between sexes in life history patterns of Sympetrum sanguineum were studied in a small pond in southern Sweden by means of exuviae and adult sampling. Emergence occurred from 4 to 28 July, and mean emergence date was 10 July for both males and females. The sex ratio at emergence (53% females) did not differ from 1:1, but significantly more females emerged during the first 5 days of the emergence period. Size of emerging individuals (immatures) decreased as season progressed and males emerged at a larger size than females. While immature males were heavier than immature females, no such difference was found in mature individuals. We suggest that the sexual differences in size and emergence patterns observed are the result of different optimisation by males and females with respect to the growth-mortality risk trade-off in the larval and adult stages.     

Developmental variation in the forest tent caterpillar: life history consequences of a threshold size for pupation

In insects, size and age at adult emergence depend on larval growth that occurs in discrete steps or instars. Understanding the mechanisms controlling stepwise larval growth and the onset of metamorphosis is essential to the study of insect life history. We examined the patterns of growth of forest tent caterpillars Malacosoma disstria to quantify variation in the number of instars that larvae undergo before pupation, to identify the mechanisms underlying variation in larval development, and to evaluate the life history consequences of this variation. All caterpillars were reared under the same conditions; at each molt, the date, the head capsule width and the mass of the freshly molted insect were recorded. Logistic regression analysis showed that a threshold size (measured either as mass or head capsule width) must be reached at the beginning of a stadium for pupation to occur at the next molt. This threshold size was higher for females than for males, and as a result, females attained a higher pupal mass than males. To achieve this larger size, females often required more instars than males, despite a higher growth ratio (size increase within an instar). Within each sex, slow growing individuals exhibited more larval instars and longer larval development time, but attained the same pupal mass as faster growing individuals. The combination of a threshold size for pupation, discrete growth steps and variation in the number of these steps can thus complicate relationships between growth rate, pupal mass and larval development time. In our study, growth ratio and number of instars were correlated with development time but not with pupal mass, and no relationship was observed between development time and pupal mass. These findings imply that, in species with variable instar number, one cannot extrapolate overall larval growth from growth during a single instar. Given the constraints of discrete larval growth, variation in instar number provides greater flexibility for insects to compensate for poor growing conditions. In this case, inferior larval growth conditions don't necessarily lead to smaller adult size.     

Sexual dimorphism in Odonata: age, size, and sex ratio at emergence

Males and females of many organisms differ in important life-history and behavioral characters. Following a recent optimization analysis of sexually dimorphic life histories, we employed an odonate-like parameter set to identify patterns of life history and behavior to be expected in an odonate population. The default parameter magnitudes generated a smaller body size and shorter development time for males than for females, which resulted in a male-biased sex ratio. Whether population growth was density dependent or density independent, and whether development time was fixed or flexible had major impacts on life-history features. The model generated five general predictions for odonate systems. (1) For species with fixed development times, males and females should differ more in activity level, growth and mortality rates than for species with flexible life cycles. (2) In species with fixed development times, populations at high latitude or high altitude should be more active, emerge and reproduce at smaller size and have a more male-biased sex ratio than low latitude and low altitude populations. (3) In density-dependent populations, with density dependence mediated by activity-dependent mortality, higher predation rates should increase activity levels and reduce development time in species with flexible development times. (4) For species with flexible development times, in strongly density-dependent populations with density dependence mediated by mortality, activity levels should decrease and development times should increase at high prey abundance. (5) Males should be larger at emergence relative to females, and the sex ratio at emergence should be more female-biased in territorial than in non-territorial species. Existing empirical evidence concerning these predictions is generally sparse and equivocal; focused tests are clearly needed.     

PROXIMATE MECHANISMS OF SEXUAL SELECTION IN WOOD FROGS

Observations and several types of field experiments on the mating behavior of wood frogs have revealed the proximate mechanisms for a size-related reproductive advantage in both males and females. For females, larger individuals produce larger clutches; for males, larger individuals can better remain clasped to females when contested by rival males and can better depose males clasped to other females. No results obtained support of the existence of mate choice in either males or females. Males were estimated to be 4.74 times as variable as females in the number of zygotes produced per individual per season; however, much of the variation in male RS resulted from a male-biased sex ratio at the breeding site rather than from sexual selection. After taking sex ratio effects into consideration, males were estimated to be only 1.63 times as variable as females. Patterns of variation in RS in males and females are associated with numerous sex-specific differences in life history and morphology. Life history differences include differential growth rates, ages at sexual maturity, and rates of mortality. Interpretation of how the body size dimorphism (females larger than males) in this species relates to sexual selection is consistent with information on how similar variations in body size influence RS for each sex, and how males and females differ in the functional relationship between body size and RS. Average RS increases more with body size in females than in males. Although body size directly influences RS for females, the possibility exists that, for males, other anatomical features correlated with body size more directly affect RS. Preliminary evidence suggests that sexual selection influences male arm length and that the male body size : RS relationship results as an incidental correlation.     

Sexual segregation in ungulates: a comparative test of three hypotheses

In most social ungulate species, males are larger than females and the sexes live in separate groups outside the breeding season. It is important for our understanding of the evolution of sociality to find out why sexual segregation is so widespread not only in ungulates but also in other mammals. Sexual body size dimorphism was proposed as a central factor in the evolution of sexual segregation in ungulates. We tested three hypotheses put forward to explain sexual segregation: the predation-risk, the forage-selection, and the activity budget hypothesis. We included in our analyses ungulate species ranging from non-dimorphic to extremely dimorphic in body size. We observed oryx, zebra, bighorn sheep and ibex in the field and relied on literature data for 31 additional species. The predation-risk hypothesis predicts that females will use relatively predator-safe habitats, while males are predicted to use habitats with higher predation risk but better food quality. Out of 24 studies on different species of ungulates, females and their offspring chose poorer quality but safer habitat in only eight cases. The forage-selection hypothesis predicts that females would select habitat based on food quality, while males should prefer high forage biomass. In fact, females selected higher quality food in only six out of 18 studies where males and females segregated, in eight studies there was no difference in forage quality and in four studies males were in better quality habitat. The activity budget hypothesis predicts that with increasing dimorphism in body size males and females will increasingly differ in the time spent in different activities. Differences in activity budgets would make it difficult for males and females to stay in mixed-sex groups due to increased costs of synchrony to maintain group cohesion. The predictions of the activity budget hypothesis were confirmed in most cases (22 out of 23 studies). The heavier males were compared to females, the more time females spent foraging compared to males. The bigger the dimorphism in body mass, the more males spent time walking compared to females. Lactating females spent more time foraging than did non-lactating females or males. Whether species were mainly bulk or intermediate feeders did not affect sexual differences in time spent foraging. We conclude that sexual differences in activity budgets are most likely driving sexual segregation and that sexual differences in predation risk or forage selection are additive factors.     

Nitrogen deposition and herbivory affect biomass production and allocation in an annual plant

If environmental conditions vary, plasticity in life-history traits is predicted. A recent model indicates that males and females should differ in life-history traits, because sexes differ in optimal attributes depending on species ecology. In this study we test the impact of two biotic factors in combination (presence/absence of predators and low/high food level) on gender specific life-history traits in the damselfly Coenagrion puella (Odonata). Results show that predator presence and low food density decreased activity in both sexes. Additionally, individuals with less food grew more slowly, emerged later, remained smaller and had a higher mortality. At low food densities, however, and in contrast to former investigations, individuals from treatments with predator presence were the same size or larger than individuals without predators. Gender had a strong impact on larval activity and life-history traits and sexes differed in development. Females were less active and took longer to complete development, but emerged at a larger size, weight and fat content. This study highlights the importance of gender specific approaches in life-history research.     

Sex-dependent effects of sibling cannibalism on life history traits of the ladybird beetle Harmonia axyridis (Coleoptera: Coccinellidae)

To investigate the sex-dependent effects of sibling cannibalism on variations in life history traits, I analysed body size, weight and instar interval in relation to the occurrence of sibling cannibalism in the ladybird beetle Harmonia axyridis. Sibling cannibalism at the time of hatching significantly affected the body size and weight of adults. There was a 2.32% and 1.05% increase in the body size of males and females, respectively, and a 3.55% increase and a 2.30% decrease in their respective body weights. Sibling cannibalism also significantly shortened the total and larval instar intervals, by 4.24% in males and by 1.22% in females, mainly due to shortening of the first instar. These results suggest that the effects of sibling cannibalism on life history traits are sex-differentiated and are greater in males than in females. A simulation of aphid density indicated that shortening the instar interval affected larval survival; the aphid density when the larvae completed development was 39.71% and 10.52% larger for cannibalistic males and females, respectively, than for non-cannibals. These results suggest that sibling cannibalism promotes more rapid development and larger adult size, although the effect was more pronounced in males than in females. Faster development may be adaptive for resource tracking, and the large adult size may increase fecundity in females and mating success in males through female mate choice, both resulting in an increase in the fitness of cannibals.  © 2002 The Linnean Society of London, Biological Journal of the Linnean Society , 2002, 76 , 349–360.     

Sexual differences in life-history traits in the butterfly Lycaena tityrus: a comparison between direct and diapause development

During direct development the butterfly Lycaena tityrus was previously found to display sex-related reaction norms in response to temperature. Based on selection for protandry in males and fecundity selection for larger females, males favoured early emergence over large size, leading to a dramatic weight loss at higher temperatures, whereas females maintained similar weights throughout. Because males were able to avoid a weight reduction relative to females in spite of their shorter development at lower temperatures, sexual size dimorphism existed at higher temperatures only. In the present paper we compare sexual differences in life-history traits in L. tityrus between direct and diapause development at 25 °C. We demonstrate that, regardless of developmental pathway, protandry persisted and relative sexual size dimorphism, with females being larger, remained unchanged. Although diapausing individuals were less time-constrained, allowing them to grow to considerably higher final weights in both sexes, males were not able to reduce their weight loss relative to females. This is explained by the pressure to gain a developmental advantage solely during post-diapause development, whereas direct developing males may spread the burden over the whole larval period. Our results highlight the importance of considering sexual differences in selective pressures, which may influence central life-history traits in manifold ways.     

Larval development of Diploptera punctata reared alone and in groups

Larvae of the cockroach Diploptera punctata were reared in isolation, in pairs, or in groups of 8–10. Duration of larval development, age at each ecdysis, weights at birth and ecdyses, and adult head-capsule width were measured. Duration of larval development was longer and adult size was larger in isolated animals than in animals reared in pairs and groups. The effect of isolation on development was more pronounced in males. All females had 4 larval instars, whereas males had 3 or 4 instars. The proportion of males with 4 larval instars was higher among animals reared in isolation. There was no difference in the duration of larval development or adult size between pair- and group-reared animals. The sex of animals in the group did not affect adult size or the duration of larval development. Males which underwent 3 or 4 larval instars had different schedules of moulting. Rates of growth of males of both instar types reared in isolation and pairs were similar. Greater adult weight of isolated animals and 4-instar-type males was a result of their longer duration of larval development. Both a higher rate of growth and longer duration of larval development contribute to the larger adult size of females than males.     

Effects of larval diet on development rates and reproductive maturation of male and female Mediterranean fruit flies

Abstract Mediterranean fruit fly Ceratitis capitata (Diptera: Tephritidae) larvae are capable of developing in one of many hosts that may vary greatly in quality. We hypothesized that they will respond to the larval environment in a manner beneficial to their subsequent reproductive performance. Accordingly, we investigated the effects of various larval diets (varying in the amount of protein and sugar they contain) on the size, development time, nutritional status and reproductive maturation (ovarian development and onset of sexual behaviour) of females and males. We found that flies which undergo larval development in artificial host fruit that contain sugar and protein ('protein-fed') were larger, developed faster and emerged with more nutritional reserves than flies that were protein-deprived as larvae. Protein-fed males, regardless of their size, became sexually active before males that developed in hosts with no protein. Protein-fed females produced more mature eggs than protein-deprived ones. Moreover, protein-fed females tended to copulate sooner than females that developed in hosts with no protein. In addition, regardless of female larval diet, females with more mature eggs tended to copulate sooner than females with less mature eggs. In light of these results, the importance of the larval environment for adult reproductive success is discussed.     

Sexual variations in food quality and gastrointestinal features of sika deer (Cervus nippon) in Japan during winter: implications for feeding strategy

We assessed sexual variation in food quality and gut macrostructure in adult male and pregnant female sika deer, Cervus nippon (Temminck, 1838), in Japan during winter. These variations might have important implications relative to sexual differences in habitat use, forage acquisition, and digestive strategy. According to the sexual dimorphism-body size hypothesis the larger males would feed on poorer forage and have heavier stomach contents and heavier intestine contents and longer intestines than smaller females. However, the food quality in rumen contents of males was higher than, or at least similar with, that of pregnant females. In correspondence to food quality, the relative weights of stomach contents and intestines with contents, the relative lengths of intestines to the lengths of body and total intestines in pregnant females were similar to adult males. The relative weights of omasum and abomasum tissues in pregnant females were greater than in males. Our findings suggest sexual differences in feeding strategy in sika deer in Japan during winter. To meet greater nutritional demands of high metabolic rate and gestation, pregnant females seemed to maintain a greater volume of digesta in guts and had more stomach tissues than expected by the sexual dimorphism-body size hypothesis to compensate for poorer forage quality.     

Environmental effects on sexual size dimorphism of a seed-feeding beetle

Sexual size dimorphism is widespread in animals but varies considerably among species and among populations within species. Much of this variation is assumed to be due to variance in selection on males versus females. However, environmental variables could affect the development of females and males differently, generating variation in dimorphism. Here we use a factorial experimental design to simultaneously examine the effects of rearing host and temperature on sexual dimorphism of the seed beetle, Callosobruchus maculatus. We found that the sexes differed in phenotypic plasticity of body size in response to rearing temperature but not rearing host, creating substantial temperature-induced variation in sexual dimorphism; females were larger than males at all temperatures, but the degree of this dimorphism was smallest at the lowest temperature. This change in dimorphism was due to a gender difference in the effect of temperature on growth rate and not due to sexual differences in plasticity of development time. Furthermore, the sex ratio (proportion males) decreased with decreasing temperature and became female-biased at the lowest temperature. This suggests that the temperature-induced change in dimorphism is potentially due to a change in non-random larval mortality of males versus females. This most important implication of this study is that rearing temperature can generate considerable intraspecific variation in the degree of sexual size dimorphism, though most studies assume that dimorphism varies little within species. Future studies should focus on whether sexual differences in phenotypic plasticity of body size are a consequence of adaptive canalization of one sex against environmental variation in temperature or whether they simply reflect a consequence of non-adaptive developmental differences between males and females.     

Egg laying preferences and larval performance in Dermestes maculatus

Dermestes maculatus DeGeer (Coleoptera: Dermestidae) is both a pest of dried stored products and, through its colonization of carrion, a forensically important species. However, little is known about the consequences of female oviposition site preferences on larval growth and development. To examine this, non‐virgin female beetles were offered a choice of food resources that had been aged to various extents to explore the adaptive nature of female oviposition preferences. Dermestes maculatus females consistently preferred to oviposit on muscle in contrast to either fat or bone marrow. Constraining larvae onto one of the three resource types confirmed that larvae grew faster and eclosed into larger adults when fed on muscle than when fed on either fat or bone marrow. In addition, the degree of sexual dimorphism was also related to food resource, with the greatest extent of size dimorphism (females larger than males) being evident on the preferred muscle resource. This conforms to the view that intraspecific variation in sexual size dimorphism is driven by intersexual differences in phenotypic plasticity, with females being able to reach greater size than males when conditions are good. The results indicate that D. maculatus female oviposition preferences are adaptive in that adult oviposition choice can enhance offspring fitness and so broadly conforms to the oviposition preference‐larval performance hypothesis as noted in a number of phytophagous insects.