Mate-Guarding as a Male Reproductive Tactic in <Emphasis Type="Italic">Propithecus verreauxi</Emphasis>

Sexual selection theory predicts that in group-living mammals, male reproductive tactics can lead to high reproductive skew in favor of dominant individuals. In sifakas (Propithecus verreauxi), a group-living primate with extremely seasonal reproduction, male reproductive success is highly skewed because dominant males sire almost all offspring despite a tendency toward an even adult group sex ratio. To understand the underlying behavioral mechanism resulting in this rank-related reproductive skew in male sifakas, we studied mate-guarding as a potential reproductive tactic. Behavioral observations of dominant males and adult females in combination with hormonal determination of timing of female receptivity in 9 groups at Kirindy Forest revealed that dominant males spent more time in proximity to females when they were receptive and were responsible for the maintenance of this proximity. Results also indicated that monopolization of receptive females was facilitated by both estrous asynchrony within groups and by the ability of dominant males to obtain olfactory cues as to the timing of female receptivity. Although dominant males engaging in mate-guarding are expected to experience various costs, there was no evidence for decreased foraging behavior and only a trend toward increased aggression between dominant and subordinate non-natal males within groups. Our results are in accordance with the hypothesis that dominant males use mate-guarding to monopolize receptive females and that it is one proximate mechanism that contributes to the high reproductive skew observed within the population of male sifakas at Kirindy.     

Male Sexual Signals and Female Choice in Drosophila grimshawi (Diptera: Drosophilidae)

Sexual selection and sexual signaling have been prominent topics in recent behavioral studies, but limited data have led to controversy regarding these topics. For example, the Hawaiian Drosophila are often cited as examples in which female choice has resulted in the evolution of elaborate male courtship signals, but relatively few data exist to test these claims adequately. We studied D. grimshawi, a lek-forming Hawaiian Drosophila, to determine whether there was evidence for female choice without male competition and to elucidate the possible cues females use to discriminate. Male mating success was found to be nonrandom and males that courted females intensely and deposited many pheromone-containing streaks on the substrate were the most successful. Hence, multiple cues seem to be involved in male mating success in this species. Some males performed only one display, however, and may represent an alternate male mating tactic. The protein content of the adult male diet significantly influenced the level of pheromone streak deposition, and thus, foraging environment may affect the outcome of sexual selection.     

An Essential Role of the Arginine Vasotocin System in Mate-Guarding Behaviors in Triadic Relationships of Medaka Fish (Oryzias latipes)

To increase individual male fitness, males of various species remain near a (potential) mating partner and repel their rivals (mate-guarding). Mate-guarding is assumed to be mediated by two different types of motivation: sexual motivation toward the opposite sex and competitive motivation toward the same sex. The genetic/molecular mechanisms underlying how mate presence affects male competitive motivation in a triadic relationship has remained largely unknown. Here we showed that male medaka fish prominently exhibit mate-guarding behavior. The presence of a female robustly triggers male-male competition for the female in a triadic relationship (2 males and 1 female). The male-male competition resulted in one male occupying a dominant position near the female while interfering with the other male''s approach of the female. Paternity testing revealed that the dominant male had a significantly higher mating success rate than the other male in a triadic relationship. We next generated medaka mutants of arginine-vasotocin (avt) and its receptors (V1a1, V1a2) and revealed that two genes, avt and V1a2, are required for normal mate-guarding behavior. In addition, behavioral analysis of courtship behaviors in a dyadic relationship and aggressive behaviors within a male group revealed that avt mutant males displayed decreased sexual motivation but showed normal aggression. In contrast, heterozygote V1a2 mutant males displayed decreased aggression, but normal mate-guarding and courtship behavior. Thus, impaired mate-guarding in avt and V1a2 homozygote mutants may be due to the loss of sexual motivation toward the opposite sex, and not to the loss of competitive motivation toward rival males. The different behavioral phenotypes between avt, V1a2 heterozygote, and V1a2 homozygote mutants suggest that there are redundant systems to activate V1a2 and that endogenous ligands activating the receptor may differ according to the social context.     

Male Choice and Male-male Competition in Idotea baitica (Crustacea,Isopoda)

Sexual selection in mate-guarding Crustacea may involve several processes: male choice, male-male competition, and female choice. To evaluate the relative importance of the different processes in mate choice of the aquatic isopod I. baitica we studied 1) the mate-choice criteria of males, 2) effects of sex ratio on the outcome of the mating contest, and 3) the role of size in male-male interactions. When given a choice between a small and a large female, males most often chose the one that matured earlier for parturial ecdysis. Maturity was a more important choice criterion than female size, but these also correlated positively. Large males had a mating advantage in both male- and female-biased sex ratios; pairing was size-assortative only in the male-biased ratio where guarding was also longer. If an extra male was placed with a precopulatory pair, 30 % take-overs occurred, large males surpassing. Present and earlier work suggests that male size is an asset in both intra- and intersexual interactions. There is little or no direct phenotypic sexual selection on female size: sexual selection for large males presumably contributes to the evolution of sexual size dimorphism in I. baitica.     

Acoustic courtship communication inSyntomeida epilais Wlk. (Lepidoptera: Arctiidae,Ctenuchinae)

The polka-dot wasp moth,Syntomedia epilais Wlk. (Lepidoptera, Arctiidae, Ctenuchinae), engages in extensive acoustic signaling during courtship. The signals, which are produced by both sexes, consist of sexually dimorphic trains of ultrasonic clicks. Field and laboratory behavioral experiments demonstrated that (1) sound production from both the male and the female is required for mating success, (2) femaleS. epilais show a higher level of acoustic responsiveness to male acoustic signals than to female signals, and (3) males use the female acoustic signals to locate their potential mates. It is suggested that the existence of the system is made possible by a reduction in bat predation pressure, perhaps the result of extreme distastefulness combined with the distinctive acoustic image that flying ctenuchines present to foraging bats.     

Prostephanus truncatus mate choice on contact: does pheromone signalling by males affect their mating success?

Prostephanus truncatus is an economically important beetle pest of stored maize and cassava in the tropics. Male beetles signal using an aggregation pheromone that attracts both female and male beetles over large distances. Females preferentially orientate towards the pheromone signals of particular males when given a choice. The influence of pheromone signalling on courtship and mating success was investigated using pheromone biossays and mating trials in both of which a female made a choice between two males. Signalling was manipulated by exposing males to a Female Factor that inhibits pheromone production. The relative attractiveness of males to females based on pheromone bioassays was found not to influence short-range courtship behaviour or mating success.     

Lack of Evidence for Energetic Costs of Mate-Guarding in Wild Male Assamese Macaques (Macaca assamensis)

Mate-guarding is a widespread and efficient male strategy for increasing paternity success. The inability to guard entire female receptive phases or complete lack of mate-guarding has been explained by energetic constraints posed on males. The energetic costs per unit time a male can afford to suffer are thought to be lowest in year-round breeding species in marginal habitats and highest in seasonally breeding species in rich habitats. Here we test the prediction that mate-guarding is energetically costly in seasonal breeders in marginal habitats. We observed all males in one group of wild Assamese macaques (Macaca assamensis) for two 4-mo mating seasons and recorded activity and travel paths via focal animal sampling, physical condition via visual inspection, and collected feces for analysis of glucocorticoid levels. Generalized linear mixed model (GLMM) analyses did not reveal an effect of mate-guarding on time spent feeding or moving or on distance traveled, travel speed, directedness of travel, or glucocorticoid levels. This lack of mate-guarding costs was consistent with the lack of a relationship between individual time spent mate-guarding and change in physical condition over the mating season. Within the limitations of our study the results do not support the idea that seasonally breeding males in marginal habitats face energetic costs of mate-guarding. With others we suggest instead that though these costs may be found in species where strength, size, and mass predict rank, mating, and reproductive success, males may not engage in costly mate-guarding in species such as Assamese macaques where also smaller, weaker males may attain high social status via political coalitions or other routes alternative to direct contest over guarded females.     

Time and energy constraints and the evolution of sexual size dimorphism — to eat or to mate?

Summary We present an empirical test of the Ghiselin—Reiss small-male hypothesis for the evolution of sexual size dimorphism (SSD). In mating systems dominated by scramble competition, where male reproductive success is a function of encounter rate with females, small males may be favoured when food is limiting because they require lower absolute amounts of food. Given a trade-off between time and energy devoted to foraging and to mate acquisition, small males should be able to devote more time to the latter. If at the same time larger females are favoured, this mechanism will contribute to the evolution of SSD and may be the major determinant of the female-biased SSDs that characterize most animal taxa. We tested this hypothesis using the water strider,Aquarius remigis (Heteroptera: Gerridae), a scramble competitor which mates many times over a prolonged mating season and which shows female-biased SSD. Laboratory experiments demonstrated that foraging success and giving up times (GUTs) are lower for males than for females during the reproductive season and that male water striders flexibly alter their time budgets under conditions of energy limitation. Controlled feeding experiments showed that male and female longevity, female fecundity and male mating success are positively related to food availability. As predicted, male body size is negatively correlated with several indices of male fitness (longevity, number of mating attempts and mating success), while female body size is positively correlated with longevity. These results are consistent with the hypothesis that scramble competition for mates favours small males in this species and provides empirical support for the Ghiselin—Reiss small-male hypothesis.     

Pupation site and emergence time influence the mating success of bagworm females,Oiketicus kirbyi

In commercial oil palm plantations in Costa Rica, we tested the hypotheses that pupation site and emergence time affect the mating success of protogynous female bagworms,Oiketicus kirbyi (Guilding) (Lepidoptera: Psychidae). Greater proportions of female than male pupae on upper leaves of oil palms and greater proportions of mated females in the upper rather than lower crown strata support the hypothesis that selection of pupation site by female larvae influences the mating success of adults. Increasing captures of males with increasing trap height further suggest that enhanced mating success of females in tree tops may be attributed either to most effective dissemination of sex pheromone on higher sites, or to males foraging predominantly in the upper strata of oil palms. As the majority of females pupated in the middle rather than upper crown of oil palms, selection of pupation site by females may be affected by additional as yet unknown factors. Emergence of females significntly preceded emergence of males. Increasing proportions of mated females throughout the emergence seasons probably resulted from an increased ‘availability’ of males. In tropical rainforests with local variations inO. kirbyi developmental time and stage, protogyny may represent an evolutionary strategy that furthers outbreeding.     

Effects of size and sex on the courting success and foraging behaviour of Embiotoca jacksoni

To test life-history theory that body size and sex should influence how animals allocate time to foraging versus reproductive activities, we measured the effects of size and sex on courting success and foraging behaviour of black surfperch Embiotoca jacksoni off Santa Catalina Island, southern California. Observations of focal fish were made while snorkelling, during which the length of each fish (estimated to the nearest cm), total duration of courting encounters and foraging rates were recorded. We made observations during and outside the mating season. Courtship occurred only between pairs and its duration increased with the size of both the male and female. Although males would court females that were smaller or larger than themselves, pairs that were closely matched in size had long courting sessions, whereas those that differed considerably in size courted only briefly. Small fish foraged more than larger fish, both during and outside the mating season. Males and females foraged at similar rates outside of the mating season, but during the mating season males reduced their foraging rates to less than half that seen outside of the mating season, whereas females continued to forage at the same rate. This decrease in foraging rate of males during the mating season was seen in all sizes of males but was proportionally greatest in the largest males. These observations indicate that males trade off time spent on foraging for time spent courting during the mating season, whereas females do not.     

Female impersonation as an alternative reproductive strategy in giant cuttlefish

Out of all the animals, cephalopods possess an unrivalled ability to change their shape and body patterns. Our observations of giant cuttlefish (Sepia apama) suggest this ability has allowed them to evolve alternative mating strategies in which males can switch between the appearance of a female and that of a male in order to foil the guarding attempts of larger males. At a mass breeding aggregation in South Australia, we repeatedly observed single small males accompanying mating pairs. While doing so, the small male assumed the body shape and patterns of a female. Such males were never attacked by the larger mate-guarding male. On more than 20 occasions, when the larger male was distracted by another male intruder, these small males, previously indistinguishable from a female, were observed to change body pattern and behaviour to that of a male in mating display. These small males then attempted to mate with the female, often with success. This potential for dynamic sexual mimicry may have played a part in driving the evolution of the remarkable powers of colour and shape transformation which characterize the cephalopods.     

Female receptivity and male mate-guarding in the jewel spiderGasteracantha minax thorell (Araneidae)

In field populations, several male jewel spidersGasteracantha minax Thorell (Araneidae) may be found at the periphery of the orb web of a female, indicating that males may compete for fertilisation success. Laboratory experiments revealed that virgin femaleG. minax readily remate shortly after their first mating. However, they appear to enter a refractory period between 1 and 24 h postmating and respond aggressively to courting males. Males that have mated with a female initially defend her from rival males but cease to do so after the onset of the refractory period. These data can be interpreted within the context of mate-guarding and sperm competition. There is marked size dimorphism in this species, which may be the result of selection for protandry. In contrast with other orb-weaving spiders, this selection pressure does not seem to be counterbalanced by selection for larger male size through either sexual cannibalism or male-male competition.     

the effect of pair formation on diel calling patterns in two cricket species,Gryllus veletis andGryllodes sigillatus (Orthoptera: Gryllidae)

After pair formation, male crickets should reduce calling to minimize the risks to males of attracting predators and/or rivals. We tested this hypothesis in two cricket species, one in which males exhibit a high mating propensity (Gryllus veletis)and another in which the mating propensity of males is constrained by the manufacture of elaborate, bipartite spermatophores (Gryllodes sigillatus).Calling durations of male G. veletisdeclined precipitously after the introduction of females but remained unchanged in G. sigillatus.We attribute the asymmetric effect of female proximity on male calling to differences in the mating propensity of males of the two species. Male G. veletisabstain from further calling in favor of repeated matings with the same female. Male G. sigillatusprobably lack the opportunity to mate repeatedly with the same female and, thus, resume calling shortly after mating to increase their probability of attracting additional mates.     

Variation in Song Rate during the Breeding Cycle of the Chaffinch,Fringilla coelebs

The variation in song rate during the breeding season was studied in two individually marked chaffinch Fringilla coelebs populations. We gathered data to investigate especially the recently presented mate-guarding hypothesis. The active singing has been supposed to function as a form of mate guarding during the female's fertile period by announcing the high status of the male and preventing extra-pair copulations by neighbouring males. There was no clear dawn chorus in the chaffinch, i.e. a peak in the song rate before sunrise. Male chaffinches continued to sing after mating, but the song rate dropped significantly. In contrast to the mate-guarding hypothesis the song rate was lower during the fertile period of the female than during pre-mating and incubation. Thus, the males do not announce the fertility status of their mates or their own quality and status by active singing. The song does not function as a form of mate guarding in the chaffinch. One function of the song of the chaffinch is mate attraction: singing activity was highest before pair formation in early spring and decreased after mating but increased again if the male lost his mate later in the breeding season.     

Vibratory Communication in the Jumping Spider Phidippus clarus: Substrate‐borne Courtship Signals are Important for Male Mating Success

Recently, work has shown that multimodal communication is common throughout the animal kingdom but the function of multimodal signals is still poorly understood. Phidippus clarus are jumping spiders in which males produce multimodal (visual and vibrational) signals in both male–male (aggressive) and male–female (courtship) contexts. The P. clarus mating system is complex, with sex ratios and the level of male competition changing over the course of the breeding season. Vibrational signal components have been shown to function in male aggressive contests but their role in courtship has not been investigated. Here, we performed an experiment to test the role of vibrational signaling in courtship by observing mating success for males that were experimentally muted. We show that vibratory courtship signals, and in particular signaling rate, is an important component of mating success and potentially a target of female choice. While the ability to produce vibratory signals significantly increased mating success, some muted males were still able to successfully mate. In these trials, signaling rate also predicted mating success suggesting that redundant signal components may compensate for errors and perturbations in signal transmission or that vibratory signals function to enhance the efficacy of visual signals.     

Laboratory adaptation reduces female mating resistance in the sweet potato weevil

Selection for genetic adaptation might occur whenever an animal colony is maintained in the laboratory. The laboratory adaptation of behavior such as foraging, dispersal ability, and mating competitiveness often causes difficulties in the maintenance of biological control agents and other beneficial organisms used in procedures such as the sterile insect technique (SIT). Sweet potato weevil, Cylas formicarius (Summers) (Coleoptera: Brentidae), is an important pest in sub‐tropical and tropical regions. An eradication program targeting C. formicarius using SIT was initiated in Japan with weevils being mass‐reared for 95 generations to obtain sufficient sterile males. The mass‐reared strain of C. formicarius exhibits weaker female resistance to male mating attempts compared with the wild strain. This could affect the success of SIT programs because mating persistence of mass‐reared males might be expected to decrease in response to weak female resistance. We show that high success of sperm transfer to mass‐reared females was due to weak female resistance to male mating attempts. However, the mating behavior of mass‐reared males did not change. In C. formicarius, the trait of male persistence to mate was not correlated with the female resistance traits. Our results suggest that mass‐rearing conditions do not have negative effects on the mating ability of the sterile males of this species, and thus that the current mass‐rearing procedures are suitable for production of sterile males for the weevil eradication program.     

The role of body size in mating success of Sphenarium purpurascens in Central Mexico

1. The effect of body size on the assortative mating and reproductive behaviour of the univoltine grasshopper Sphenarium purpurascens (Charpentier) was studied in Central Mexico. 2. Assortative mating by size was observed in the field. Evidence of positive assortative mating in relation to body size was found in laboratory experiments. Female fecundity and male success in contests were also correlated with body size. 3. Larger females had a higher number of eggs per pod. Larger males usually won fights and were able to take over females from other males, and to resist takeovers by other males while guarding. 4. Individuals of both sexes were observed copulating with more than one sexual partner in the field, suggesting polygamy. Male–male contests determined access to females, and males exhibited a postcopulatory prolonged mate-guarding behaviour lasting up to 18 days. 5. In a 2-year study, sex ratio was male-biased at the beginning of the reproductive season and decreased to 1:1 by the end of the season, suggesting that the population is protandrous. 6. The results of this study indicate that assortative mating results from male–male competition and female availability, and suggests that body size is a potential target of natural and sexual selection.     

同种雄性竞争对手的存在对星豹蛛雄蛛求偶和交配行为的影响

越来越多的研究发现,雄性产生精子(精液)也需付出代价。雄性除了依据配偶质量和竞争对手的竞争强度适应性调整生殖投入外,雄性在求偶和交配行为上也相应产生适应性反应,求偶和交配行为具有可塑性。目前雄性求偶和交配行为可塑性研究主要集中于雌性多次交配的类群中,在雌性单次交配的类群中研究甚少。以雌蛛一生只交配一次而雄蛛可多次交配的星豹蛛为研究对象,比较:(1)前一雄性拖丝上信息物质对后续雄蛛求偶和交配行为的影响,(2)雌雄不同性比对雄蛛求偶和交配行为的影响。研究结果表明,星豹蛛前一雄蛛拖丝上的信息物质对后续雄蛛求偶潜伏期、求偶持续时间和交配持续时间都没有显著影响,但前一雄蛛拖丝上的信息物质对后续雄蛛求偶强度有显著抑制作用。同时,性比对星豹蛛雄蛛求偶和交配行为都没有显著影响。可见,星豹蛛雄蛛对同种雄性拖丝上的化学信息可产生求偶行为的适应性调整,而对性比不产生适应性反应。     

Male foraging avoidance in female feeding territories in a harem polygynous cichlid in Lake Tanganyika

We studied foraging site partitioning between the sexes in Neolamprologus tetracanthus, a shrimp-eating Tanganyikan cichlid with harem-polygyny. Females maintained small territories against heterospecific food competitors within large territories of males, foraging exclusively at the inner side of their own territories (foraging areas). Males fed as frequently as females in their own territories, but mostly outside female foraging areas, although they frequently entered female territories and repelled food competitors from the territories. Soon after removal of the resident females, however, harem males, as well as many food competitors, invaded the vacant territories and intensively devoured prey of female foraging areas. This indicates that although female foraging areas appear to contain more food than outside the areas, harem males refrained from foraging there when the resident females were present. We suggest that harem males will attempt to keep female foraging areas in good condition, whereby they may get females to reside in male territories and/or promote female gonadal maturation.     

Within‐Species Mate Preferences Do Not Contribute to the Maintenance of Sexually Monomorphic Mating Signals in Green Lacewings

We know much less about the evolutionary forces and constraints that maintain similar mating displays in females and males than we do about sexually dimorphic mating displays. Both female and male green lacewings have sexually monomorphic vibrational mating signals and are equally choosy against heterospecific mating signals. This similarity in between‐species sex roles may explain a large part of the presence of species‐specific female signals in these species, but does not necessarily predict why female and male signals are similar. We tested for within‐species sex‐specific similarities in mate preferences in Chrysoperla lucasina that may contribute to the maintenance of sexually monomorphic mating signals in this species. We found weak preferences and low levels of discrimination for signals with varying fine‐scale temporal features (volley duration, period, and volley‐duty cycle). The longer signals that both sexes produced in response to playback were sexually monomorphic, but some females and most males also produced shorter signals with significantly reduced volley durations and periods. Notably, all of these signals had indistinguishable volley‐duty cycles, the ratio of volley duration to volley period. We propose that mating signals in C. lucasina are maintained in both sexes because of similar between‐species mate preferences, but the sexually monomorphic mating signals cannot be attributed to significant within‐species mate preferences. What differences are present in within‐species sex roles may be resolved by a male‐biased signal polymorphism, in long and short signals that are hypothesized to have distinct functions during mate calling and courtship.