Longevity cost of reproduction for males but no longevity cost of mating or courtship for females in the male-dimorphic dung beetle Onthophagus binodis

Life history theory predicts a trade-off between current and future reproduction. Despite a wealth of research on the cost of reproduction for females, there have been very few studies that have looked at the cost of reproduction for males. Longevity is closely related to the opportunity for future reproduction, and thus decreased longevity in response to current reproductive effort has been used as a measure of the cost of reproduction. Here we examine the cost of reproduction for males and females in the dung beetle Onthophagus binodis. Like many onthophagines, O. binodis exhibit dimorphic male morphology; major males develop a large pronotal horn while minor males remain hornless. Alternative morphologies are associated with alternative reproductive tactics. Thus, we ask whether major and minor males pay different costs of reproduction. We found that in contrast to previous work on Diptera, mating is not costly in terms of reduced longevity for female dung beetles. Despite a longevity cost of reproduction for males, we found no evidence for differential longevity costs associated with alternative reproductive tactics.     

Flight dimorphism is related to survival,reproduction and mating success in the leaf beetle Oreina cacaliae

1. Alternative life histories may be maintained in populations due to variation in the costs and benefits of the underlying strategies. In this study, potential costs of dispersal by flight were investigated as an alternative life‐history strategy in the mountain‐living chrysomelid beetle Oreina cacaliae. 2. In this species, previous mark–recapture studies showed a dispersal dimorphism in both males and females. While a fraction of the population engages in flight in autumn and spring (in the following referred to as ‘flyers’), the other part does not fly (non‐flyers). Flyers emerge earlier than non‐flyers and feed on a spring host plant before the emergence of the main host plant. 3. In this study, the overwintering and dispersal locations were recorded over 7 years in the field, flyers from the spring host plant were collected, and morphology and lifetime reproductive output and survival of collected flyers and non‐flyers were compared. 4. A potential trade‐off between flight and life‐history traits was observed: flyers were smaller in size, lighter in body mass, had a lower lifetime fecundity and a higher mortality. 5. Mating experiments of field‐caught beetles in the laboratory showed that larger beetles had a higher (multiple) mating success, but there was no evidence for size‐assortative mating. It is hypothesized that one reason for small beetles to disperse by flight might be to escape competition for mates with larger non‐flyers. 6. The overwhelming quantity of beetles found on the spring host every year reveals that the flying strategy is successful, despite the costs and risks.     

Female mate assessment and choice behavior affect the frequency of alternative male mating tactics

Explanations for the existence of alternative male mating tacticsfocus primarily on male–male competition. Mating systems,however, are composed of interactions both within and betweenthe sexes, and the role of female behavior in shaping male matingtactics should not be overlooked. By using a dynamic state variablegame model, I examine how female mate assessment and choicebehavior affect the frequency of alternative male mating tactics.When females can accurately assess the quality of males, onlymales with high quality are likely to be chosen as mates, andthus, lower-quality males gain little fitness from courtingfemales. This leads lower-quality males to switch to an alternativemating tactic that attempts to circumvent female mate choice.In contrast, if the abilities of females to accurately assessmales are constrained by assessment costs, imperfect information,or time constraints, or if the pool of available males is smaller,then lower-quality males are increasingly chosen as mates andthey less often use alternative mating tactics. Thus, femalebehavior shapes the frequency of alternative male mating tactics.A consequence of this game between the sexes is that male behavior(i.e., increased alternative mating tactics) decreases the benefitsfemales might otherwise gain from lower assessment costs, clearersignals of male quality, more time to choose a male, and moremales from which to choose a mate.     

Single and multiple mating reduces longevity of female dumpling squid (Euprymna tasmanica)

For many species, mating is a necessary yet costly activity. The costs involved can have an important influence on the evolution of life histories and senescence. Females of many species mate multiply, and this behaviour can inflict a longevity cost. Most studies investigating the effects of multiple mating on female survival have been conducted on insects, and the effects in other taxa are largely unknown. We investigate the effects of both a single mating and a second mating on longevity in female dumpling squid (Euprymna tasmanica), a species in which both sexes mate multiply. Through comparing the longevity of virgin, once‐mated and twice‐mated females, we found that a single mating reduced female life span by 15 days on average. A second mating resulted in an additional 8 day (on average) longevity cost, despite no difference in total clutch mass, number of clutches, single egg mass or number of eggs per clutch between once‐mated and twice‐mated females. This demonstrates a cost to multiple mating which may be independent of the cost of egg production. Furthermore, total clutch mass and female life span were positively correlated, whereas female life span decreased with increasing average water temperature. The presence of an additive effect of reproduction on longevity suggests that multiple mating in cephalopods may have benefits that outweigh these costs, or that there is a conflict in optimal mating frequency between males and females.     

The rate of fertilization in male mating tactics of the polygynous grey seal

Studies using molecular markers have shown that some grey seal males may be gaining success through exhibiting alternative mating tactics. We estimated the probability of fertilization success of grey seal males exhibiting the primary tactic of female defence and one alternative tactic of mating with departing females on Sable Island, Nova Scotia, Canada, during the breeding seasons of 1997-2002. Although the fertilization rate of the primary tactic (27-43%) was greater than that of the alternative tactic (10-12%), these low rates indicate the potential fitness value of alternative mating tactics in this size-dimorphic pinniped species.     

Resource availability,mating opportunity and sexual selection intensity influence the expression of male alternative reproductive tactics

The expression of alternative reproductive tactics can be plastic and occur simultaneously depending on cues that vary spatially or temporally. For example, variation in resources and sexual selection intensity is expected to influence the pay‐off of each tactic and shape the decision of which tactic to employ. Males of the nuptial gift‐giving spider Pisaura mirabilis can adopt three tactics: offering a genuine prey gift, a ‘worthless’ non‐nutritious gift or no gift. We hypothesized that resources and/or male body condition, and mating opportunity and sexual selection intensity, vary over the course of the mating season to shape the co‐existence of alternative traits. We measured these variables in the field over two seasons, to investigate the predictions that as the mating season progresses, (i) males become more likely to employ a gift‐giving tactic, and (ii) the likelihood of switching from worthless to genuine gifts increases. Prey availability increased over the season and co‐varied with the propensity of males to employ the gift‐giving tactic, but we found no support for condition‐dependent gift giving. Males responded to an increase in female availability by increasing their mating effort (gift production). Furthermore, the frequency of genuine gift use increased with sexual selection intensity, consistent with the assumption that sperm competition intensity increases with time. Our results suggest that the frequency of alternative tactics is shaped by seasonal changes in ecological factors and sexual selection. This leads to relaxed selection for the gift‐giving tactic early in the season when females are less choosy and resources more scarce, and increased selection for genuine gifts later in the season driven by mating opportunity and risk of sperm competition.     

Alternative male mating tactics of the endemic Red Sea parrotfish Scarus ferrugineus

The present study shows that small non‐territorial terminal‐phase males of the rusty parrotfish Scarus ferrugineus are reproductively active and are comparable with initial‐phase males in behaviour, rates of participation during group‐spawning and success in streaking into pair spawning. Large territorial terminal‐phase males defend contiguous territories for several hours during the morning where they pair spawn with initial‐phase females.     

Confidence of paternity and paternal care: covariation revealed through the experimental manipulation of the mating system in the beetle Onthophagus taurus

Theoretical models of paternal care predict that facultative reductions in male care may occur under certain conditions. One important parameter that has been shown to influence the outcome of these models is a male's confidence of paternity. In this study, we tested whether the amount of care provided by horned males in the dimorphic beetle, Onthophagus taurus, varied with his confidence of paternity. Male care results in an increased weight of dung provided in the brood masses produced by the pair. Using the sterile male technique we showed that a horned male's paternity declined with the number of sneak males in the population. The relationship was nonlinear, with paternity declining most rapidly between a frequency of one and three sneaks, and stabilizing thereafter at about 50%. A horned male's paternity was directly related to the number of copulations with the female, relative to the number of copulations achieved by sneaks. Horned males were shown to reduce their care in relation to their declining paternity. Video analysis demonstrated that reductions in male care occurred through a combination of male desertion and a trade‐off between caring and paternity assurance behaviours. The number of fights with sneak males was negatively related to the amount of care provided by a horned male. These results suggest that by gauging his expected paternity through the number of fights with sneaks, a horned male is able to assess his paternity and reduce his investment accordingly. Our data thus provide strong empirical support for the proposed link between paternity and paternal care.     

Cost of reproduction in Callosobruchus maculatus: effects of mating on male longevity and the effect of male mating status on female longevity

One of the most studied life-history trade-offs is that resulting from the cost of reproduction: a trade-off arises when reproduction diverts limited resources from other life-history traits. We examine the cost of reproduction in male, and the effect of male mating status on female Callosobruchus maculatus seed beetles. Cost of reproduction for male C. maculatus was manifested as reduced longevity. There was also a positive relationship between male body size and male longevity. Females mated to males that had already copulated twice did not live as long as females mated to males that had copulated once or not at all. The third copulation of males also lasted longer than the two previous ones. We conclude that even though the cost of reproduction for males has been studied much less than that in females, there is growing evidence that male reproductive effort is more complex than has traditionally been thought.     

Wing length and asymmetry of male Tokunagayusurika akamusi chironomid midges using alternative mating tactics

Male Tokunagayusurika akamusi chironomids have alternative mating tactics.One is to search for females on vegetation (ground mating),and the other is to wait for females in an aerial swarm (swarmmating). Simultaneous sampling of ground-unpaired and ground-pairedmales and of swarm-unpaired and swarm-paired males were performed.The average wing length and right-left wing length difference(wing asymmetry) were compared between males from the four differentcategories. Swarm-unpaired males were larger than ground-unpaired ones,swarm-paired males were larger than swarm-unpaired ones, and ground-pairedmales were not larger than ground-unpaired ones. Thus, large malestended to aggregate in swarms, and larger swarming males matedmore successfully. On the other hand, small males probably enjoyedmating on the ground, especially when large males swarmed. Thewing asymmetry was not significantly different between unpairedand paired males both within and between tactics. There wasa flat or U-shaped relationship between wing length and asymmetry,underpinning the lack of a symmetrical advantage of swarmingto large males. The right-left difference was not normally distributedin four of six samples of unpaired males but, in contrast, wasnot normally distributed in only one of six samples of pairedmales. The non-normal distributions were leptokurtic and includedoutliers. Removal of the outliers improved normality, suggestingthat males with extremely asymmetric wings were not successfulin mating.     

The effects of age at mating on female life-history traits in a seed beetle

Age at first reproduction is an important component of lifehistory across taxa and can ultimately affect fitness. Becausegenetic interests of males and females over reproductive decisionscommonly differ, theory predicts that conflict may arise overthe temporal distribution of matings. To determine the potentialfor such sexual conflict, we studied the direct costs and benefitsassociated with mating at different times for females, usingseed beetles (Acanthoscelides obtectus) as a model system. Virginfemales were resistant to male mating attempts at a very earlyage but subsequently reduced their resistance. Although we foundno difference in life span or mortality rates between femalesmated early in life and those mated later, females that matedearly in life suffered a 12% reduction in lifetime fecundity.Thus, there are direct costs associated with mating early inlife for females. Yet, males mate even with newly hatched females.We suggest that these data indicate a potential for sexual conflictover the timing of first mating and that female resistance tomating, at least in part, may represent a female strategy aimedat delaying mating to a later time in life.     

Investment in survival and reproduction along a semelparity–iteroparity gradient in the Beta species complex

The Beta species complex shows a gradient of life histories from pronounced semelparity (big‐bang reproduction) to pronounced iteroparity (repeated reproduction). Models assume a trade‐off between investment in reproduction and survival. Reproductive effort is thought to increase with decreasing life span, and to be invariable in semelparous plants and susceptible to environmental conditions in iteroparous plants. These assumptions and hypotheses were verified by a greenhouse experiment testing six different life cycles at three contrasting nutrient levels. This study suggests that reproductive effort is negatively correlated with mean life span along the life‐cycle gradient. Unlike semelparous beets, reproductive effort in iteroparous beets is extremely sensitive to nutrient level. Phenotypic correlation between allocation to reproduction and allocation to survival generally appeared significantly negative in the longest‐lived iteroparous beets, nonsignificant in intermediate life histories and obviously positive in semelparous beets (no trade‐off control).     

Female access and diet affect insemination success,senescence and the cost of reproduction in the male Mexican fruit fly Anastrepha ludens

Hypotheses exploring the influence of dietary conditions on the life‐history trade‐off between survival and reproductive success are extensively tested in female insects but only rarely explored in males. The present study examines the impact of dietary quality and female access on age‐specific reproduction and survival of the male Mexican fruit fly Anastrepha ludens Loew (Diptera: Tephritidae). There is a clear cost of female access for males with access to dietary protein, measurable as a decrease in life expectancy, which is further influenced by the age when females are introduced. A protein deficient diet reduces the lifespan benefit of virginity and masks the detrimental effect of female access on male life expectancy. Dietary protein is not necessary for reproductive success, although access to protein at eclosion improves the lifetime reproductive success of males compared to when it is delayed. Overall, reproductive success diminishes as the male flies age, regardless of the dietary conditions, providing evidence for reproductive senescence in males. Delaying the males' access to a protein source fails to influence the negative effect of age on reproductive ability. Because age‐specific reproductive rates decline with age, regardless of diet, male fitness does not benefit from lifespan extension. Therefore, males can be expected to allocate available resources towards reproductive effort in favour of an extended lifespan, regardless of mate and protein availability.     

The effect of male mating history on paternal investment, fecundity and female remating in the seed beetle Callosobruchus maculatus

1. In many organisms, males provide nutrients to females via ejaculates that can influence female fecundity, longevity and mating behaviour. The effect of male mating history on male ejaculate size, female fecundity, female longevity and female remating behaviour in the seed beetle Callosobruchus maculatus was determined.
2. The quantity of ejaculate passed to females declined dramatically with successive matings. Despite the decline, a male's ability to fertilize a female fully did not appear to decline substantially until his fourth mating.
3. When females multiply mated with males of a particular mated status, the pattern of egg production was cyclic, with egg production increasing after mating. Females multiply mated to virgins had higher fecundity than females mated to non-virgins, and females mated to twice-mated males had disproportionately increased egg production late in their life.
4. Females that mated to multiple virgins, and consequently laid more eggs, experienced greater mortality than females mated only once or mated to non-virgins, suggesting that egg production is costly, and rather than ameliorating these costs, male ejaculates may increase them by allowing or stimulating females to lay more eggs.
5. Females mating with non-virgin males remated more readily than did females mated to virgins. Females given food supplements were less likely to remate than females that were nutritionally stressed, suggesting that females remate in part to obtain additional nutrients.     

Competition from large males and the alternative mating tactics of small males of dawson’s burrowing bee (Amegilla dawsoni) (apidae, apinae, anthophorini)

Males of Dawson’s burrowing bees (Amegilla dawsoni) search for virgin females at three locations: (1) open clay patches where females are emerging from underground brood cells, (2) the vegetated peripheral zone adjacent to emergence areas (through which females pass after emerging), and (3) clusters of flowering plants, which are often some distance from emergence areas. Males of Dawson’s burrowing bees exhibit a size dimorphism with large major and small minors. Major males patrol only the open emergence sites, whereas minor males may be found in all three locations. Although most females are mounted and presumably mated immediately upon emergence, some are not, and these females make up a pool of potential mates for the small males patrolling the peripheral zone and flower patches. The density of males at emergence sites and the probability of male-male aggression change over the course of a day and over the entire flight season. When the level of competition is low, some minor males hunt for mates at emergence areas, where potential mates are relatively numerous. But when the presence of many large rivals makes it unlikely that a small male can avoid being displaced from emerging females, minors make the best of a bad job by shifting to areas where majors are absent.     

Heat shock proteins mediate trade‐offs between early‐life reproduction and late survival in Drosophila melanogaster

Ageing and the resulting increased likelihood mortality are the inescapable fate of organisms because selection pressures on genes that exert their function late in life is weak, promoting the evolution of genes that enhance early‐life reproductive performance at the same time as sacrificing late survival. Heat shock proteins (HSP) are known to buffer various environmental stresses and are also involved in protein homeostasis and longevity. The characteristics of genes for HSPs (hsp) imply that they affect various life‐history traits, which in turn affect longevity; however, little is known about the effects of hsp genes on life‐history traits and their interaction with longevity. In the present study, the effects of hsp genes on multiple fitness traits, such as locomotor activity, total fecundity, early fecundity and survival time, are investigated in Drosophila melanogaster Meigen using RNA interference (RNAi). In egg‐laying females, RNAi knockdown of six hsp genes (hsp22, hsp23, hsp67Ba, hsp67Bb, hsp67Bc and hsp27‐like) does not shorten survival but rather increases it. Knockdown of five of those genes on an individual basis reduces early‐life reproduction, suggesting that several hsp genes mediate the trade‐off between early reproduction and late survival. The data indicate a positive effect of hsp genes on early reproduction and also negative effects on survival time, supporting the antagonistic pleiotropic effects predicted by the optimality theory of ageing.     

绿缘扁角叶甲生活史、交配及繁殖的观察

【目的】探明绿缘扁角叶甲Platycorynus parryi Baly年生活史、交配及繁殖情况。【方法】越冬代成虫在室内自然条件下连续繁殖,及在不同温度下,详细观察记录了成虫的交配、繁殖力和寿命。【结果】绿缘扁角叶甲是络石Trachelospermum jasminoides的重要害虫,以老熟幼虫在3~5 cm深的土层中越冬,在江西南昌一年发生1代。越冬幼虫于次年4月中旬开始陆续化蛹,4月下旬成虫开始羽化。自然条件下各虫态发育历期:卵期(11.81±0.17)d,幼虫期300~330 d,蛹期(13.62±0.13)d,成虫寿命(47.99±0.65)d。成虫羽化3~4 d后开始交配,日平均交配(3.55±0.11)次,交配持续时间最短5 min,最长可达140 min,平均交配持续时间(28.11±0.89)min,日龄大的成虫交配持续时间显著短于日龄小的成虫;相邻两次交配之间的间隔时间最短3 min,最长426 min,平均交配间隔时间(74.75±3.19)min,日龄大的成虫交配间隔时间显著短于日龄小的成虫。交配后2~3 d开始产卵,平均每雌产卵量为169粒。在22、25和28℃条件下,温度对雌虫寿命具有显著影响,随着温度升高,雌虫寿命显著缩短,而雄虫寿命和每雌产卵量却差异不显著。【结论】绿缘扁角叶甲在江西南昌一年发生1代,以老熟幼虫在土中越冬,成虫可进行多次交配,成虫日龄对其交配行为有显著影响,温度对该虫的繁殖力无显著影响。     

Determinants of male ejaculate investment in the cotton bollworm Helicoverpa armigera: mating history,female body size and male age

Many studies demonstrate that ejaculate size may be influenced by male condition, female quality and the risk or intensity of sperm competition. In the present study, the effect of male and female conditions, male mating history and female mating status on ejaculate sperm numbers in the polyandrous moth Helicoverpa armigera is examined. A large variation in ejaculate size is found and, although female body size and male age influence ejaculate size, female age and copula duration do not. Both male and female mating histories have significant effects on ejaculate sperm numbers. Males reduce ejaculate expenditure in successive matings but deliver significantly more apyrene and eupyrene sperm to nonvirgin than to virgin females.     

Evaluating the life‐history trade‐off between dispersal capability and reproduction in wing dimorphic insects: a meta‐analysis

A life‐history trade‐off exists between flight capability and reproduction in many wing dimorphic insects: a long‐winged morph is flight‐capable at the expense of reproduction, while a short‐winged morph cannot fly, is less mobile, but has greater reproductive output. Using meta‐analyses, I investigated specific questions regarding this trade‐off. The trade‐off in females was expressed primarily as a later onset of egg production and lower fecundity in long‐winged females relative to short‐winged females. Although considerably less work has been done with males, the trade‐off exists for males among traits primarily related to mate acquisition. The trade‐off can potentially be mitigated in males, as long‐winged individuals possess an advantage in traits that can offset the costs of flight capability such as a shorter development time. The strength and direction of trends differed significantly among insect orders, and there was a relationship between the strength and direction of trends with the relative flight capabilities between the morphs. I discuss how the trade‐off might be both under‐ and overestimated in the literature, especially in light of work that has examined two relevant aspects of wing dimorphic species: (1) the effect of flight‐muscle histolysis on reproductive investment; and (2) the performance of actual flight by flight‐capable individuals.