Life History Effects Upon Contest Behaviour: Age as a Predictor of Territorial Contest Dynamics in Two Populations of the Speckled Wood Butterfly, Pararge aegeria L.

Although empirical studies of life history effects upon sexually selected phenomena have largely overlooked contest behaviour, recent research suggests that territorial contest participation in butterflies may be mediated by ageing per se. Verbal and mathematical arguments predict lifetime increases in the expression of risky male reproductive behaviours, such as fighting, under a range of ecological conditions. Here we explored the relevance of ageing per se to contest dynamics in two phenologically distinct populations of the speckled wood butterfly, Pararge aegeria. We established 160 experimental like‐population contests among naïve south Swedish and Madeiran dyads, 112 of which we varied the age difference between combatants by 3–4 d. Although this age asymmetry did not influence contest outcome in either population, we found weak positive covariance between the loser's age and contest duration amongst Madeiran males. This is consistent with a slight lifetime increase in aggression because the duration of these aerial persistence contests is a sensitive measure of the losing male's level of aggressive intent. However, the size of this effect (semi‐partial correlation = 0.281) suggests age is not as strongly relevant to contest behaviour in P. aegeria as in other territorial butterflies. We discuss the ways in which ecological differences between butterfly species, particularly with respect to predation risk, may have influenced the evolution of lifetime aggressive strategies in this group.     

Residual reproductive value and male mating success: older males do better

Evolution via sexual selection has traditionally been viewed as isolated from life-history constraints. As additionally reproductive resource allocation in males is underexplored, it is rather unclear how life-history factors have shaped lifetime investment into male sexually selected traits. Against this background, we here investigate male butterfly mating success in relation to age, nutritional status, assay condition and wing damage. As predicted, based on a low residual reproductive value, older males had a considerably higher mating success than younger males. Comparisons between virgin and once-mated males suggest that this pattern is related to age per se rather than differential ratings of the resource receptive female. We found no evidence for male body size or condition being important, supporting the notion that in weaponless animals intrinsic motivation is more important for mating success than the differences in physical properties (such as body size or condition). Flight cage experiments suggest that such differences in motivation may be masked under more natural conditions, where flight performance, having a clear impact on mating success (as evidenced by wing manipulation experiments), is likely to be crucial. We conclude that the life-history perspective is a fruitful one for gaining a better understanding of the evolution of sexually selected characters and the predictions derived from contest theory do also apply to male mating success.     

Coerced first sexual intercourse and selected reproductive health outcomes among young women in KwaZulu-Natal, South Africa

Coercion may play an important role in compelling young women to engage in sexual intercourse at an early age. With a decline in age at first intercourse and increased reporting of coercive first sex, concerns of adverse reproductive health outcomes such as unintended pregnancy and sexually transmitted infections (STIs), including HIV, have become important issues, particularly in the context of a high HIV prevalence. This paper uses data collected in 2001 from the second round of a longitudinal study of 1130 sexually experienced young women in KwaZulu-Natal to investigate the relationship between coerced first intercourse and selected reproductive health outcomes and behaviours. Nearly 46% of all sexually experienced young women had reported that their first sexual encounter had been coerced. Young women who reported being coerced at first sex were significantly more likely to be Black and living in an urban area. Those who had been coerced at first sex were also more likely to report having had an STI and having experienced unintended pregnancy, than those who had not been coerced at first sex. Coercion at first sex is an important social and public health problem that has a serious impact on the reproductive health and behaviours of young women. Interventions should directly address the issue of sexual coercion by ensuring young women are aware of their reproductive rights.     

The costs of risky male behaviour: sex differences in seasonal survival in a small sexually monomorphic primate

Male excess mortality is widespread among mammals and frequently interpreted as a cost of sexually selected traits that enhance male reproductive success. Sex differences in the propensity to engage in risky behaviours are often invoked to explain the sex gap in survival. Here, we aim to isolate and quantify the survival consequences of two potentially risky male behavioural strategies in a small sexually monomorphic primate, the grey mouse lemur Microcebus murinus: (i) most females hibernate during a large part of the austral winter, whereas most males remain active and (ii) during the brief annual mating season males roam widely in search of receptive females. Using a 10-year capture-mark-recapture dataset from a population of M. murinus in Kirindy Forest, western Madagascar, we statistically modelled sex-specific seasonal survival probabilities. Surprisingly, we did not find any evidence for direct survival benefits of hibernation-winter survival did not differ between males and females. By contrast, during the breeding season males survived less well than females (sex gap: 16%). Consistent with the 'risky male behaviour' hypothesis, the period for lowered male survival was restricted to the short mating season. Thus, sex differences in survival in a promiscuous mammal can be substantial even in the absence of sexual dimorphism.     

The marginal valuation of fertility

Substantial theoretical and empirical evidence demonstrates that fertility entails economic, physiological, and demographic trade-offs. The existence of trade-offs suggests that fitness should be maximized by an intermediate level of fertility, but this hypothesis has not had much support in the human life-history literature. We suggest that the difficulty of finding intermediate optima may be a function of the way fitness is calculated. Evolutionary analyses of human behavior typically use lifetime reproductive success as their fitness criterion. This fitness measure implicitly assumes that women are indifferent to the timing of reproduction and that they are risk-neutral in their reproductive decision-making. In this paper, we offer an alternative, easily-calculated fitness measure that accounts for differences in reproductive timing and yields clear preferences in the face of risky reproductive decision-making. Using historical demographic data from a genealogically-detailed dataset from 19th century Utah, we show that this measure is highly concave with respect to reproductive effort. This result has three major implications: (1) if births are properly timed, a lower-fertility reproductive strategy can have the same fitness as a high-fertility strategy, (2) intermediate optima are far more likely using fitness measures that are strongly concave with respect to effort, (3) we expect mothers to have strong investment preferences with respect to the risk inherent in reproduction.     

Experimentally increased badge size increases male competition and reduces male parental care in the collared flycatcher

Experimental enlargement of sexually selected traits that are energetically cheap to produce is expected to reveal costs resulting from increased risk of predation or social competition. Given a trade-off between sexually selected traits and life history traits such costs may be expected to affect not only the males themselves but also their offspring. In this study I manipulated the size of the forehead patch, a sexually selected trait that functions as a badge of status in male collared flycatchersFicedula albicollis). First, I found that a male''s likelihood to establish a breeding territory with respect to his original badge size was affected by the treatment such that old males (older than or equal to two years) with relatively small original badges enjoyed an increased likelihood of establishing a breeding territory while young males (yearlings) suffered a reduced likelihood of establishment when their badges were enlarged as compared to unchanged. Second, young males with enlarged badges that were able to establish a territory fed their nestlings less in relation to their females compared to the control males. However, the females adjusted their parental effort to such an extent that no significant differences were observed in total feeding rate nor in reproductive success between the two groups of males. These results suggest that experimentally enlarged badge size in the collared flycatcher may result in increased male competition and that males have to trade their effort spent in male contest against their parental effort.     

Male eastern bluebirds trade future ornamentation for current reproductive investment

Life-history theory proposes that organisms must trade-off investment in current and future reproduction. Production of ornamental display is an important component of reproductive effort that has rarely been considered in tests of allocation trade-offs. Male eastern bluebirds (Sialia sialis) display brilliant ultraviolet-blue plumage that is correlated with mate acquisition and male competitive ability. To investigate trade-offs between current reproductive effort and the future expression of a sexually selected ornament, we manipulated the parental effort of males by changing their brood sizes. We found that parents provisioned experimentally enlarged broods more often than reduced broods. As predicted by life-history theory, the change in parental effort had a significant effect on the relative plumage ornamentation of males in the subsequent year: males with reduced broods significantly increased in plumage brightness. Moreover, this change in plumage coloration had a direct effect on the timing of breeding in the following season: males that displayed brighter plumage in the year following the manipulation mated with females that initiated egg laying earlier in the season. These data indicate that male bluebirds must trade-off conserving energy for production of future ornamentation versus expending energy for current reproduction.     

Song predicts immunocompetence in male European starlings (Sturnus vulgaris)

According to the immunocompetence handicap hypothesis, sexually selected characteristics predict immune function and this relationship is mediated by testosterone. In the present study, we investigated whether bird song could predict immunity in European starlings (Sturnus vulgaris). We recorded the singing and reproductive behaviours of 16 adult male starlings in an outdoor aviary and then assessed their cell-mediated and humoral immunity in vivo. The males were observed in groups of four for 2 h each day over a 4-day period. For each male, the number of songs produced was recorded and the average song-bout length was computed. Next, cell-mediated and humoral immunity were assessed via cutaneous swelling responses to the T-cell mitogen phytohaemagglutinin and antibody responses to a novel antigen, keyhole limpet haemocyanin. Song rate and song-bout length were positively correlated with cell-mediated and humoral immunity, respectively. Additionally, a negative relationship between plasma testosterone concentration and antibody response was observed. These data demonstrate that male starling song can be used as a predictor of immunocompetence, with more robust singers exhibiting enhanced immunity. Whether this relationship is mediated directly by testosterone requires further investigation.     

Correlation between Male Social Status,Testosterone Levels,and Parasitism in a Dimorphic Polygynous Mammal

Life history trade-offs have often been assumed to be the consequence of restrictions in the availability of critical resources such as energy and nutrients, which necessitate the differential allocation of resources to costly traits. Here, we examined endocrine (testosterone) and health (parasite burdens) parameters in territorial and non-territorial New Zealand fur seal males. We documented intra-sexual differences in sexual behaviours, testosterone levels, and parasitism that suggest a trade-off exists between reproductive success and physical health, particularly susceptibility to helminths and acanthocephalans, in males displaying different mating tactics (i.e., territorial and non-territorial tactics). Levels of testosterone were higher in territorial males and correlated positively with reproductive effort (i.e., intra- and inter-sexual interactions). However, these territorial males also exhibited high levels of parasitic infection, which may impair survival in the long-term. Our study, while limited in sample size, provides preliminary evidence for a link between male mating tactics, testosterone levels and parasite loads, and potential effects on reproductive success and life history that should be explored further.     

Cancer susceptibility and reproductive trade-offs: a model of the evolution of cancer defences

The factors influencing cancer susceptibility and why it varies across species are major open questions in the field of cancer biology. One underexplored source of variation in cancer susceptibility may arise from trade-offs between reproductive competitiveness (e.g. sexually selected traits, earlier reproduction and higher fertility) and cancer defence. We build a model that contrasts the probabilistic onset of cancer with other, extrinsic causes of mortality and use it to predict that intense reproductive competition will lower cancer defences and increase cancer incidence. We explore the trade-off between cancer defences and intraspecific competition across different extrinsic mortality conditions and different levels of trade-off intensity, and find the largest effect of competition on cancer in species where low extrinsic mortality combines with strong trade-offs. In such species, selection to delay cancer and selection to outcompete conspecifics are both strong, and the latter conflicts with the former. We discuss evidence for the assumed trade-off between reproductive competitiveness and cancer susceptibility. Sexually selected traits such as ornaments or large body size require high levels of cell proliferation and appear to be associated with greater cancer susceptibility. Similar associations exist for female traits such as continuous egg-laying in domestic hens and earlier reproductive maturity. Trade-offs between reproduction and cancer defences may be instantiated by a variety of mechanisms, including higher levels of growth factors and hormones, less efficient cell-cycle control and less DNA repair, or simply a larger number of cell divisions (relevant when reproductive success requires large body size or rapid reproductive cycles). These mechanisms can affect intra- and interspecific variation in cancer susceptibility arising from rapid cell proliferation during reproductive maturation, intrasexual competition and reproduction.     

The evolution of sexually selected traits and antagonistic androgen expression in actinopterygiian fishes

Many sexually selected traits in male fishes are controlled by testosterone. Directional selection for male ornaments could theoretically increase male testosterone levels over evolutionary timescales, and when genetically correlated, female testosterone levels as well. Because of the negative fitness consequences of high testosterone, it is plausible that female choice for sexually selected traits in males results in decreased female reproductive fitness. I used comparative analysis to examine the association between male peak testosterone expression and sexually selected ornaments. I also tested for genetic correlation between male and female androgen levels. The presence of sexually selected traits in males was significantly correlated with increased peak androgen levels in males as well as females, and female testosterone levels were significantly correlated with male peak testosterone titers, although the slope was only marginally <1. This suggests that selection to decouple high male and female testosterone levels is either weak or otherwise ineffective.     

Parental and Mating Effort: Is There Necessarily a Trade‐Off?

One of the common assumptions in the study of the evolution of parental care is that trade-offs exist between parental investment and other fitness-related traits. In general, this body of work follows the traditional definition that parental investment (in the current offspring) decreases that individual's ability to invest in future reproduction ( Trivers 1972 ). However, examination of the empirical evidence shows that assuming a trade-off between parental and mating effort is not always appropriate. This overemphasis on a trade-off between mating and parental effort has arisen in part because of an oversimplification of female reproductive strategies, a failure to consider interactions between the sexes, and a tendency to consider behaviours as unifunctional, thereby ignoring the more complex relationship between mating and parental effort in many species. Here, we first examine the empirical evidence for trade-offs between mating and parental effort in males and females to ask when trade-offs occur and what pattern they take. By highlighting a number of exemplar species, we then explore how the presence or absence of trade-offs relates to mate choice and sexual selection in both sexes. Finally, we highlight the importance of considering individual variation, which has been particularly overlooked in examinations of female investment, and how preferences in one sex may influence the existence and our interpretation of apparent trade-offs in the other sex.     

Male‐mediated prenatal loss: Functions and mechanisms

Sexually selected infanticide has been the subject of intense empirical and theoretical study for decades; a related phenomenon, male‐mediated prenatal loss, has received much less attention in evolutionary studies. Male‐mediated prenatal loss occurs when inseminated or pregnant females terminate reproductive effort following exposure to a nonsire male, either through implantation failure or pregnancy termination. Male‐mediated prenatal loss encompasses two sub‐phenomena: sexually selected feticide and the Bruce effect. In this review, we provide a framework that explains the relationship between feticide and the Bruce effect and describes what is known about the proximate and ultimate mechanisms involved in each. Using a simple model, we demonstrate that male‐mediated prenatal loss can provide greater reproductive benefits to males than infanticide. We therefore suggest that, compared to infanticide, male‐mediated prenatal loss may be more prevalent in mammalian species and may have played a greater role in their social evolution than has previously been documented.     

Climate change and condition of herring (Clupea harengus) explain long-term trends in extent of skipped reproduction

It is commonly assumed that iteroparous fish, once mature, normally reproduce in all consecutive seasons. Recent work has suggested, however, that in Norwegian spring-spawning herring—a population that undertakes extensive spawning migrations—almost one in two adults may skip their second spawning migration. Why should herring not return to spawn the year after first spawning, but instead wait an extra year? For herring, participation in distant, energetically costly, and risky spawning migrations will only pay off in terms of fitness if individuals are sufficiently large, and in sufficient condition, to both successfully migrate and spawn. Changes in the environment and individual condition should therefore affect the likelihood of skipped spawning. This paper describes long-term changes in the extent to which the second reproductive season is skipped in this herring population. These are shown to be linked to the size and condition of herring as first-time spawners, and to climatic factors possibly related to food availability. The findings corroborate the hypothesis that skipped reproduction results from trade-offs between current and future reproduction, growth and survival.     

Chest Color and Social Status in Male Geladas (<Emphasis Type="Italic">Theropithecus gelada</Emphasis>)

Conspicuous colored patches on animals often serve as sexually selected signals that advertise male quality. Such colored traits facilitate assessment of risks associated with a specific contest or benefits associated with a specific mate choice. Here, we investigate whether a colored patch of skin on the chests of male geladas (Theropithecus gelada) is a sexually selected signal. Specifically, we examine the relationship between color (redness), social status (a proxy for reproductive success), and age. We use observational data from known individuals from a population of wild geladas living in Ethiopia. We digitally quantified chest color using a previously-validated method for measuring color under field conditions. Results from this study are consistent with the hypothesis that redness is a quality signal in males. Baseline color correlates with status even when controlling for age. Indeed, males with redder chests were members of “better” groups: 1) leader males—the only males with reproductive access to females—had the reddest chests, and 2) within leader males, males with large units (>6 females) had redder chests than males with small units. At present, we are unable to address whether male chest color is directed at potential rivals or mates. Nevertheless, our data support the hypothesis that quality signals should prevail in large, fluid groups, where it is unlikely that individuals recognize all other group members. If individual recognition is limited in gelada society, this would favor the evolution of alternative means of assessment for making reproductive decisions.     

Experimental evidence for adaptive personalities in a wild passerine bird

Individuals of the same species differ consistently in risky actions. Such ‘animal personality’ variation is intriguing because behavioural flexibility is often assumed to be the norm. Recent theory predicts that between-individual differences in propensity to take risks should evolve if individuals differ in future fitness expectations: individuals with high long-term fitness expectations (i.e. that have much to lose) should behave consistently more cautious than individuals with lower expectations. Consequently, any manipulation of future fitness expectations should result in within-individual changes in risky behaviour in the direction predicted by this adaptive theory. We tested this prediction and confirmed experimentally that individuals indeed adjust their ‘exploration behaviour’, a proxy for risk-taking behaviour, to their future fitness expectations. We show for wild great tits (Parus major) that individuals with experimentally decreased survival probability become faster explorers (i.e. increase risk-taking behaviour) compared to individuals with increased survival probability. We also show, using quantitative genetics approaches, that non-genetic effects (i.e. permanent environment effects) underpin adaptive personality variation in this species. This study thereby confirms a key prediction of adaptive personality theory based on life-history trade-offs, and implies that selection may indeed favour the evolution of personalities in situations where individuals differ in future fitness expectations.     

Pushing Wasps to Work: Decentralized Aggression Induces Increased Activity in the Paper Wasp <Emphasis Type="Italic">Polistes versicolor</Emphasis>

When helpers from cooperative breeding animals have some expectation of direct reproduction, there is potential for conflict over how much aid they should provide to the colony. For example, if food is shared among all colony members, then higher levels of foraging by a helper would be desirable for the colony as a whole. However, because foraging is risky and physiologically costly, hopeful reproductive helpers could avoid foraging. Evidence suggests that this work-conflict could be resolved if helpers are aggressively coerced by their nestmates to provide aid. Here, we showed that in the primitively eusocial paper wasp Polistes versicolor, colony food starvation leads to an increasing in aggression that results in an increasing activity level (including foraging). We propose that aggression affects forage levels because (i) attacks from nestmates are directed toward known foragers rather than non-foragers; and (ii) resting wasps generally respond to aggression by becoming more active while already active wasps generally respond by switching the task they were doing. In P. versicolor, direct reproduction is an option for helpers. It means that they can be considered as hopeful reproductive individuals seeking to avoid performing risky behaviours, like foraging. In this sense, decentralized aggression from nestmates could be a coercive mechanism to force wasps performing undesirable tasks, while simultaneously enhancing the performance of a variety of other tasks.     

Effects of a food supplementation experiment on reproductive investment and a post-mating sexually selected trait in magpies Pica pica

Food availability is an important factor affecting breeding success in birds. Food supplementation experiments in birds have in general focused on the effects on reproductive success in terms of female investment (laying date, clutch size, egg size), however, it is also known that the estimation of mate quality based on sexually selected signals influences female reproductive investment. In the particular case of magpies, females use nest size, a post-mating sexually selected signal, to assess male's likelihood to invest in reproduction, and accordingly adjust reproductive investment (clutch size). Then, the possible effects of food supplementation on female reproductive investment could be mediated by other variables related to parental quality, such as nest size in magpies. In the present study, we explore if higher food availability in a magpie territory affected both male sexually selected traits (i.e. nest size) and female reproductive investment (laying date, egg size, clutch size). We performed a food supplementation experiment in which we experimentally increased food availability in several magpie territories, keeping others as controls. In food-supplemented territories, males built significantly larger nests and females significantly increased egg size by 4.1% compared to control females. Results suggest that the continuous provisioning of protein rich food allowed magpie females to increase egg size. However, laying date and clutch size did not differ between control and food-supplemented magpie pairs. Food availability also affected the relationship between female reproductive investment and nest size. In control territories, females decreased their egg size in response to a larger nest, whereas a tendency for the opposite relationship was revealed in food-supplemented territories. We discuss the possibility that magpie females adopt different strategies for reproductive investment according to food availability.     

A possible link between parasite defence and residual reproduction

Life-history theory centres around trade-offs between current and future reproduction, but we have little understanding of how such trade-offs are mediated. We supplementary fed Ural owls (Strix uralensis) during the nestling period and quantified parents' current and future life-history components as well as their physiological health by monitoring haematocrit, leucocyte profile, intra- and extracellular blood parasites. Feeding led to reduced parental effort but did not improve offspring viability, male parasite defence, or parental survival. Intracellular leucocytozoan infection was reduced in fed females which lasted to the following year's reproductive season (carry-over effect), when fed females also laid larger and earlier clutches. Leucocytozoon infection therefore may mediate the life-history trade-off between current and residual reproduction in this species.     

SELECTION FOR ALTERNATIVE MALE REPRODUCTIVE TACTICS ALTERS INTRALOCUS SEXUAL CONFLICT

Intralocus sexual conflict (IASC) arises when fitness optima for a shared trait differ between the sexes; such conflict may help maintain genetic variation within populations. Sex‐limited expression of sexually antagonistic traits may help resolve the conflict, but the extent of this resolution remains a subject of debate. In species with alternative male reproductive tactics, unresolved conflict should manifest more in a more sexually dimorphic male phenotype. We tested this prediction in the bulb mite (Rhizoglyphus robini), a species in which aggressive fighters coexist with benign scramblers. To do this, we established replicated lines in which we increased the proportion of each of the alternative male morphs using artificial selection. After approximately 40 generations, the proportion of fighters and scramblers stabilized at >0.9 in fighter‐ and scrambler‐selected lines, respectively. We then measured several female fitness components. As predicted by IASC theory, female fecundity and longevity were lower in lines selected for fighters and higher in lines selected for scramblers. This finding indicates that sexually selected phenotypes are associated with an ontogenetic conflict that is not easily resolved. Furthermore, we suggest that IASC may be an important mechanism contributing to the maintenance of genetic variation in the expression of alternative reproductive tactics.