Males,but not females,perform strategic mate searching movements between host plants in a leaf beetle with scramble competition polygyny

Mate searching is assumed to be performed mostly by males, but when females benefit from multiple mating or are under risk of failing to mate, they may also perform mate searching. This is especially important in scramble competition polygynies, in which mate searching is the main mechanism of mate competition. Typically, more mobile individuals are expected to achieve higher mating success because mobility increases their probability of finding mates. If we assume individual movements are mainly explained by mate searching in scramble competition polygynies, we can investigate searching strategies by asking when individuals should leave their location and where they should go. We hypothesize that individuals will leave their locations when mating opportunities are scarce and will seek spatially close sites with better mating opportunities. We tested these hypotheses for males and females of Leptinotarsa undecimlineata, a leaf beetle with scramble competition polygyny in which both sexes are promiscuous. Individuals mate and feed exclusively on Solanum plants, and thus, individual movements can be described as switches between plants. Females were less likely than males to leave isolated plants, and both males and females moved preferentially to neighboring plants. Males were more likely to leave when the local number of females was low, and the number of males was high. They moved to plants with more females, a behavior consistent with a mate searching strategy. Females were more likely to move to plants with fewer males and many females, a behavior consistent with male harassment avoidance. Strategic movement is widely considered in foraging context, but seldom in a mate searching context. Considering that selection to minimize searching costs, maximize mating success, and minimize harassment may be ubiquitous in nature, we argue that strategic movements by mate searching individuals are likely to occur in many species.     

Sex roles and mutual mate choice matter during mate sampling

The roles of females and males in mating competition and mate choice have lately proven more variable, between and within species, than previously thought. In nature, mating competition occurs during mate search and is expected to be regulated by the numbers of potential mates and same-sex competitors. Here, we present the first study to test how a temporal change in sex roles affects mating competition and mate choice during mate sampling. Our model system (the marine fish Gobiusculus flavescens) is uniquely suitable because of its change in sex roles, from conventional to reversed, over the breeding season. As predicted from sex role theory, courtship was typically initiated by males and terminated by females early in the breeding season. The opposite pattern was observed late in the season, at which time several females often simultaneously courted the same male. Mate-searching females visited more males early than late in the breeding season. Our study shows that mutual mate choice and mating competition can have profound effects on female and male behavior. Future work needs to consider the dynamic nature of mating competition and mate choice if we aim to fully understand sexual selection in the wild.     

Complex mate searching in the satin bowerbird Ptilonorhynchus violaceus

Mate-choice studies typically focus on male traits affecting female mating decisions, but few studies seek to identify the behavioral rules females use when searching for mates. Current models suggest that females may either directly compare a set of males ("pooled comparison") or compare each male to an internal standard ("sequential-search rule") when judging the suitability of potential mates. Models also differ in other specific aspects, such as the predicted number of sampling bouts initiated and the tendency of females to return to males after previous visits. We monitored 63 female satin bowerbirds, Ptilonorhynchus violaceus, during mate sampling to reconstruct their search patterns. We found that females typically sampled several males and returned to the most attractive male for mating: a behavior consistent with the pooled-comparison tactic. Females, however, varied in the number of males sampled; some visited only one male before mating. We found that this variation can be explained by differences among females in the number of mates, the date mate searching is initiated, and long-term experience with males. Further, females were observed to initiate two distinct sampling bouts, with the rejection of most of their potential mates occurring before the start of the second sampling bout. This suggests that the choices of potential mates are narrowed prior to the second sampling bout and that the later visits may function to reconsider preliminary decisions made during the first sampling bout or to resolve decisions concerning the remaining potential mates. Our results indicate that mate searching is a complex process in which females use multiple sampling bouts to find suitable mates and in which several different factors influence their search behavior.     

Dynamic mate-searching tactic allows female satin bowerbirds Ptilonorhynchus violaceus to reduce searching

Females can maximize the benefits of mate choice by finding high-quality mates while using search tactics that limit the costs of searching for mates. Mate-searching models indicate that specific search tactics would best optimize this trade-off under different conditions. These models do not, however, consider that females may use information from previous years to improve mate searching and reduce search costs in subsequent years. We followed female satin bowerbirds Ptilonorhynchus violaceus during mate searching and reconstructed their search patterns. We found that females who chose very attractive males typically mated with the same male in the following year, resulting in these females sampling fewer males than those who switched mates. In contrast, females who mated with less attractive males typically rejected their previous mates and searched longer for more attractive mates in the following mating season. A potential cost to mate searching is suggested by the observed increase in the likelihood of force-copulation attempts from marauding males with increased searching. Our results suggest that by using past experiences to adjust their search tactics, females may obtain high-quality mates while limiting search costs. These results emphasize the need to consider historical effects in studies of sexual selection, especially for long-lived species with stable display sites.     

Female mate choice and mating costs in the polyandrous butterflyPieris napi (Lepidoptera: Pieridae)

Males of the green-veined white butterfly (Pieris napi L.) transfer large ejaculates that represent on average 15% of their body mass when mating for a first time. Shortly after mating a male is able to transfer only a small ejaculate when mating a second time. Male ejaculate production plays a crucial role in the mating system ofP. napi because females use male-derived nutrients for egg production and somatic maintenance. Here we study how timing of female rematings and copulation duration are influenced by the mating history of their mates and, also, study if females exert mate choice to minimize their mating costs. Mating with a recently mated male increased female mating costs by increasing time in copula and mating frequency. Virgin females that mated with virgin males remated after an average of 6 days, whereas virgin females that mated with recently mated males remated after an average of 2 days. Moreover, copulations involving recently mated males lasted on average almost 7 h, whereas copulations involving virgin males lasted on average 2 h. Recently mated males were eager to remate, in spite of the fact that the size of the ejaculate they transfer is small and that they remain in copula for a long time. Hence it seems that males are more successful in the sexual conflict over mating decisions and that females do not minimize mating costs by choosing to mate preferentially with virgin males.     

Do males choose their mates in the lekking moth Achroia grisella? Influence of female body mass and male reproductive status on male mate choice

Lekking males aggregate to attract females and contribute solely to egg fertilization, without any further parental care. Evolutionary theory therefore predicts them to be nonchoosy toward their mates, because any lost mating opportunities would outweigh the benefits associated with such preferences. Nevertheless, due to time costs, the production of energetically costly sexual displays, and potential sperm limitation, the mating effort of lekking males is often considerable. These factors, combined with the fact that many females of varying quality are likely to visit leks, could favor the evolution of male mate preferences. Here, we show that males of the lekking lesser wax moth, Achroia grisella, were indeed more likely to mate with heavier females in choice experiments, even at their virgin mating (i.e., when their reproductive resources have not yet been depleted by previous matings). This differential female mating success could not be attributed to female behavior as heavy and light females showed similar motivation to mate (i.e., latency to approach the males) and time to copulate. Males seem to benefit from mating with heavier females, as fecundity positively correlated with female mass. This new empirical evidence shows that male mate choice may have been underestimated in lekking species.     

视觉和触角在栗山天牛雄虫近距离搜寻配偶行为中的作用

栗山天牛Massicus raddei (Blessig)是中国栎树类树种的主要蛀干害虫, 在吉林省和辽宁省主要危害蒙古栎Quercus mongolica和辽东栎Quercus liaotungensis的主干, 给当地造成了重大的经济损失和生态灾害。为了明确是否有信息化学物质参与到雄性成虫的配偶搜寻和识别过程中, 采用行为观测笼, 分别对栗山天牛雄虫的视觉和嗅觉作用进行了研究。结果显示： 致盲后的雄虫能进行正常的配偶搜寻和交尾, 而且正常的雄虫对只有视觉刺激而无化学刺激的蜡封雌虫尸体毫无反应, 说明视觉对雄虫搜寻配偶所起的作用很小。切除7节以上鞭节的雄虫交尾次数大大下降, 切除全部9节鞭节的雄虫完全不能实现交尾行为, 而切除全部鞭节的雌虫能正常交尾, 说明雄虫的触角特别是其端部7节鞭节在识别配偶并成功交尾方面起着重要的作用, 并且可能由其感受雌虫所释放的信息化学物质。     

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.     

Impact of the Timing of Male Emergence on Mating Capacity of Males in Trichogramma evanescens Westwood

In species with highly structured population, such as Trichogramma spp., mating occurs mostly at emergence on the patch and early emerging males have an advantage, as they are present when the first females emerge. However, early emergence of male could also enable males to mature sexually before the emergence of females or enhance their capacity to induce higher receptivity in females. We measured time of emergence of male and female Trichogramma evanescens Westwood (Hymenoptera: Trichogrammatidae) from hosts that were parasitized within a 30 min period. We also measured the effect of male age on their capacity to mate, and the ability of inseminated females to produce daughters. To verify if early emerging males induced higher receptivity in females, we observed the females mate choice between males of different ages. Our results indicated that the mean time of emergence between males and females was 29 min for individuals that develop in 9 days and 10 min for individuals that develop in 10 days. The protandry observed in this species could be the result of either a faster development of males or a male first strategy by the ovipositing female. In T. evanescens, protandry does not permit males to mature sexually nor to induce higher receptivity in females. The main advantage of protandry for males thus appears to be early access to females as they emerge.     

Is Male Cohabitation Costly for Females of the Spider Stegodyphus lineatus (Eresidae)?

Reproductive interests of females and males can vary over a number of issues, including the number of matings and the occurrence and duration of mate guarding and cohabitation. The mating system of the spider Stegodyphus lineatus (Eresidae) is characterized by an exceptional sexual conflict where males induce females to remate by committing infanticide. Females experience high costs because of this male strategy, while males benefit. During the mating season, males tend to stay with females for several days. We examined whether this male strategy of cohabitation is also disadvantageous for females of S. lineatus. A field experiment revealed that male presence in a female's nest negatively affected her body condition, whereas cohabiting males gained weight because they fed on prey caught in webs of females. As a response, females did not renew their webs when males were present. Thus, females with a cohabiting male experienced the combined cost of prey loss and loss of time available for foraging. These costs are expected to increase with every additional cohabiting male. Mated females behaved more aggressively toward males than did virgin females. This behavior may be interpreted as an adaptive response to reduce mating costs.     

Energetic costs of mate guarding behavior in male stream-dwelling isopods

In the stream-dwelling isopod Lirceus fontinalis, males and females engage in a precopulatory mate guarding phase prior to mating. We examined the energetic costs of mate guarding behavior in males by separately assaying glycogen and lipid content at different time increments following mating. We found that males that had recently mated possessed reduced glycogen reserves and that these reserves were fully replenished within 36 h. Conversely, we found that male lipid reserves were unaffected by time since mating. We concluded that precopulatory mate guarding behavior is energetically costly to males and that glycogen is the energy source utilized to pay that cost. We also examined whether food deprivation during the mate guarding phase affected male energy reserves (glycogen) at the end of that phase. We found that males that were held in the laboratory and starved during mate guarding possessed reduced glycogen at the termination of the phase when compared to fed males. This reduced quantity was equivalent to the glycogen reserves of recently mated males collected from the field. We propose that food deprivation during the mate guarding phase explains the reduction in glycogen reserves at the termination of that phase. We discuss these results with reference to patterns of refuge use behavior during the mate guarding phase.     

Reproductive costs of mating with a sibling male: sperm depletion and multiple mating in Cephalonomia hyalinipennis

Polygynous parasitoid males may be limited by the amount of sperm they can transmit to females, which in turn may become sperm limited. In this study, I tested the effect of male mating history on copula duration, female fecundity, and offspring sex ratio, and the likelihood that females will have multiple mates, in the gregarious parasitoid Cephalonomia hyalinipennis Ashmead (Hymenoptera: Bethylidae: Epyrinae), a likely candidate for sperm depletion due to its local mate competition system. Males were eager to mate with the seven females presented in rapid succession. Copula duration did not differ with male mating history, but latency before a first mating was significantly longer than before consecutive matings. Male mating history had no bearing on female fecundity (number of offspring), but significantly influenced offspring sex ratio. The last female to mate with a given male produced significantly more male offspring than the first one, and eventually became sperm depleted. In contrast, the offspring sex ratio of first‐mated females was female biased, denoting a high degree of sex allocation control. Once‐mated females, whether sperm‐depleted or not, accepted a second mating after a period of oviposition. Sperm‐depleted females resumed production of fertilized eggs after a second mating. Young, recently mated females also accepted a second mating, but extended in‐copula courtship was observed. Carrying out multiple matings in this species thus seems to reduce the cost of being constrained to produce only haploid males after accepting copulation with a sperm‐depleted male. I discuss the reproductive fitness costs that females experience when mating solely with their sibling males and the reproductive fitness gain of males that persist in mating, even when almost sperm‐depleted. Behavioural observations support the hypothesis that females monitor their sperm stock. It is concluded that C. hyalinipennis is a species with a partial local mating system.     

Vagabond males and sedentary females: spatial ecology and mating system of the speckled rattlesnake (Crotalus mitchellii)

We used radiotelemetric data and behavioural observations to characterize seasonal (mating versus post‐mating seasons) and sexual variation in movement patterns, as well as to examine some of the ecological factors contributing to the evolution of the mating system in a venomous predator from the Mojave Desert of North America, the speckled rattlesnake, Crotalus mitchellii. Mating occurs in spring from late April to early June, shortly after emergence from hibernation, when snakes are predictably aggregated around the dens. Males and females travelled further per unit time in the mating season compared to the post‐mating season. Males also travelled longer distances per unit time than females in the mating and post‐mating seasons, and males with larger home ranges during the mating season had more potential mating partners. The results obtained suggest that males actively locate females during the mating season, and that the drastic increase in distance travelled by males during the mating season may be caused by strong male–male competition for access to females, probably because of the limited availability of sexually receptive females. Furthermore, males fight for access to females, and males of larger size are more likely to acquire females. Therefore, sexual selection apparently acts on two different male phenotypic traits: investment in mate‐searching activities and male body size. The present study demonstrates that combining quantitative spatial analyses and behavioural observations in an explicit temporal context can significantly advance our understanding of the ecology and evolution of organismal mating systems. © 2011 The Linnean Society of London, Biological Journal of the Linnean Society, 2011, 103 , 681–695.     

Effects of parasitic infection on mate sampling by female wild turkeys (Meleagris gallopavo): should infected females be more or less choosy?

Investigations of parasite-mediated sexual selection have concentratedon the effects of parasites on males. Differences in femalesusceptibility to parasitic infection may also cause variationin reproductive behavior. I propose two alternative hypothesesto explain how infected females may alter their mate samplingbehavior. In the first hypothesis, infected females sample fewerprospective mates because chronic parasitic infection imposesenergetic costs that limit the time and calories that a femalecan expend in mate searching. A novel alternative hypothesisis that females recognize their own susceptibility to infectionand thus invest more time searching for a male phenotype thatindicates he offers genes complementary to her genome. In recombination,these good genes would allow her offspring to better resistparasites despite their mother's susceptibility. I examinedthe mate sampling behavior of experimentally infected wild turkeyhens when presented with an array of males, and compared themto control hens. Infected females did not invest more time assessingindividuals, did not wait longer to choose a male, nor werethey less likely to solicit during the trial. They did differfrom control females in that they visited more males beforesoliciting copulation and exhibited different preference functionsfor snood length. These results suggest that females are notso energetically restricted by latent coccidia infection thatthey must hurry to find a mate. Instead, it appears that infectedfemales assess a larger set of males as prospective mates, perhapsto increase the opportunity to obtain complementary genes forparasite resistance.     

Plasticity in male mating behavior modulates female life history in fruit flies

In many species, intense male-male competition for the opportunity to sire offspring has led to the evolution of selfish reproductive traits that are harmful to the females they mate with. In the fruit fly, Drosophila melanogaster, males modulate their reproductive behavior based on the perceived intensity of competition in their premating environment. Specifically, males housed with other males subsequently transfer a larger ejaculate during a longer mating compared to males housed alone. Although the potential fitness benefits to males from such plasticity are clear, its effects on females are mostly unknown. Hence, we tested the long-term consequences to females from mating with males with distinct social experiences. First, we verified that competitive experience influences male mating behavior and found that males housed with rivals subsequently have shorter mating latencies and longer mating durations. Then, we exposed females every other day for 20 days to males that were either housed alone or with rivals, and subsequently measured their fitness. We found that females mated to males housed with rivals produce more offspring early in life but fewer offspring later in life and have shorter lifespans but similar intrinsic population growth rates. These results indicate that plasticity in male mating behavior can influence female life histories by altering females’ relative allocation to early versus late investment in reproduction and survival.     

Male size does not affect the strength of male mate choice for high-quality females in Drosophila melanogaster

Theory predicts that the strength of male mate choice should vary depending on male quality when higher-quality males receive greater fitness benefits from being choosy. This pattern extends to differences in male body size, with larger males often having stronger pre- and post-copulatory preferences than smaller males. We sought to determine whether large males and small males differ in the strength (or direction) of their preference for large, high-fecundity females using the fruit fly, Drosophila melanogaster. We measured male courtship preferences and mating duration to show that male body size had no impact on the strength of male mate choice; all males, regardless of their size, had equally strong preferences for large females. To understand the selective pressures shaping male mate choice in males of different sizes, we also measured the fitness benefits associated with preferring large females for both large and small males. Male body size did not affect the benefits that males received: large and small males were equally successful at mating with large females, received the same direct fitness benefits from mating with large females, and showed similar competitive fertilization success with large females. These findings provide insight into why the strength of male mate choice was not affected by male body size in this system. Our study highlights the importance of evaluating the benefits and costs of male mate choice across multiple males to predict when differences in male mate choice should occur.     

Males Benefit from Mating with Outbred Females in Drosophila littoralis: Male Choice for Female Genetic Quality?

The evolution and expression of mate choice behaviour in either sex depends on the sex‐specific combination of mating costs, benefits of choice and constraints on choice. If the benefits of choice are larger for one sex, we would expect that sex to be choosier, assuming that the mating costs and constraints on choice are equal between sexes. Because deliberate inbreeding is a powerful genetic method for experimental manipulation of the quality of study organisms, we tested the effects of both male and female inbreeding on egg and offspring production in Drosophila littoralis. Female inbreeding significantly reduced offspring production (mostly due to lower egg‐to‐adult viability), whereas male inbreeding did not affect offspring production (despite a slight effect of paternal inbreeding on egg‐to‐adult viability). As inbreeding depressed female quality more than male quality, the benefits of mate choice were larger for males than for females. In mate choice experiments, inbreeding did not affect male mating success (measured as a probability to be accepted as a mate in a large group), suggesting that females did not discriminate among inbred and outbred males. In contrast, female mating success was affected by inbreeding, with outbred females having higher mating success than inbred females. This result was not explained by lower activity of inbred females. Our results show that D. littoralis males benefit from mating with outbred females of high genetic quality and suggest adaptive male mate choice for female genetic quality in this species. Thus, patterns of mating success in mate choice trials mirrored the benefits of choice: the sex that benefited more from choice (i.e. males) was more choosy.     

Male and Female Soldier Beetles Relax Choice for Mate Quality Across Daily Courtship Periods

Mutual mate choice occurs when members of each sex will reject some potential mates in efforts to encounter better prospects later. The decision to reject may represent the interaction between mate preferences, mate availability, and temporal constraints. Theory predicts that mutual mate choice will favor relaxed choosiness as mate availability and time for courtship decline. We explored mutual mate choice in the soldier beetle, Chauliognathus pennsylvanicus (Cantharidae), where courtship consisted of males attempting to secure evasive females. We employed field observations, laboratory experiments, and stochastic simulations to investigate the decline in choosiness over the daily courtship period, during which individuals can mate at most once. We found that reproductive success of males and females increases with mate size and mating frequency. Females biased copulations toward larger mates by attempting to evade suitors, while males biased copulations by releasing the smaller females they capture. However, late in the day males and females may increase reproductive success by accepting rather than rejecting lower quality mates to maintain high mating rates. Stochastic simulations indicated that reproductive success, the product of mating frequency and mean mate size, was maximized in males and females by incrementally reducing mate standards across daily courtship periods. In the field, large males who rejected small females early in the daily courtship period rarely did so later. Large females used less effective evasive maneuvers later in the courtship period, resulting in copulations with progressively smaller males. These results support models of mutual mate choice that predict that individuals of high quality will maximize reproductive success by relaxing choosiness as the courtship period wanes.     

Male mate choice in the chameleon grasshopper (Kosciuscola tristis)

In many species, males can increase their fitness by mating with the highest quality females. Female quality can be indicated by cues, such as body size, age and mating status. In the alpine grasshopper Kosciuscola tristis, males can be found riding on subadult females early in the season, and as the season progresses, males engage in fights over ovipositing females. These observations suggest that males may be competing for females that are either unmated (early season) or sperm‐depleted (late season). We thus hypothesised that male K. tristis may be choosy in relation to female mating status, and specifically, we predicted that males prefer females that are unmated. We conducted behavioural experiments in which males were given the choice of two females, one mated and one unmated. Contrary to our prediction, males did not mate preferentially with unmated females. However, copulation duration with unmated females was, on average, 24 times the length of copulation with mated females. While female K. tristis can reject mates, we did not observe any evidence of overt female choice during our trials. Females may gain additional benefits from mating multiply and may therefore not readily reject males. While our experiment cannot definitively disentangle female from male control over copulation duration, we suggest that males choose to invest more time in copula with unmated females, perhaps for paternity assurance, and that male mate assessment occurs during copulation rather than beforehand.