Male Mating Strategies Research Articles

Functional morphology of the visual system and mating strategies in bumblebees (Hymenoptera, Apidae, <italic>Bombus</italic>)

Bumblebee mating behaviour intrigued Darwin over 150 years ago and has elicited much interest since. Despite our increasing knowledge about male mating strategies, we still know little about the signals and cues involved, the sender-receiver relationship and the functional adaptations of the involved sensory systems. Although several authors have noticed a close relationship between the mating system and male-specific elaboration of the visual system, studies on the functional adaptations have yet to be conducted. We investigated the functional morphology of the compound eyes in 11 species of Bombus from various subgenera in detail. Of these, four species (Bombus confusus, Bombus melaleucus, Bombus mendax, Bombus niveatus) have enlarged male eyes, characterized by a higher number of ommatidia compared with worker eyes and a frontal zone with enlarged facets, which is likely to be associated with improved spatial resolution and contrast sensitivity. In these species, perching mating strategies are found. In contrast, males of species that patrol scent routes (Bombus hortorum, Bombus lapidarius, Bombus pascuorum, Bombus pratorum, Bombus soroeensis, Bombus terrestris, Bombus wurflenii) show no distinct eye adaptations, and their eye morphology closely resembles that of the workers. A phylogenetic analysis of male eye structure indicates that enlarged males eyes may have been the ancestral state in Bombus.

Can males detect the strength of sperm competition and presence of genital plugs during mate choice?

Female reproductive status can have strong selective effects on male mating strategies due to the threat of sperm competition, which may explain why males prefer virgin over mated females. However, in mating systems with female multiple mating rates and mating plugs, males should not only respond to the risk but also to the level of sperm competition and should be sensitive to the interference from mating plugs. In the orb-web spider Argiope keyserlingi, females possess paired sperm-storage organs facilitating separate sperm storage from different males. Males are limited to 2 copulations due to mutilation of their paired genitals (pedipalps). By conducting binary choice experiments, we tested whether males of different mating status can distinguish between females that mated with 1 or 2 males and whether single-mated males discriminate between single-mated females with matched or unmatched virgin genital openings. Furthermore, we investigated whether males adapt their mating strategies to the intensity of sperm competition by providing males with varying qualities of females in their immediate vicinity. Our results demonstrated that males are sensitive to the level of sperm competition and preferred single-mated females over double-mated females. However, they failed to identify single-mated females whose virgin genital opening matched their unused pedipalp, which is required due to their fixed ipsilateral insemination. Surprisingly, males never mated twice with the same female independently of the quality of surrounding females. This suggests that the benefits from searching and mating with a different female are greater than the benefits of monopolizing the female by mating twice with her.

Functional morphology of the visual system and mating strategies in bumblebees (Hymenoptera, Apidae,Bombus)

Bumblebee mating behaviour intrigued Darwin over 150 years ago and has elicited much interest since. Despite our increasing knowledge about male mating strategies, we still know little about the signals and cues involved, the sender-receiver relationship and the functional adaptations of the involved sensory systems. Although several authors have noticed a close relationship between the mating system and male-specific elaboration of the visual system, studies on the functional adaptations have yet to be conducted. We investigated the functional morphology of the compound eyes in 11 species of Bombus from various subgenera in detail. Of these, four species (Bombus confusus, Bombus melaleucus, Bombus mendax, Bombus niveatus) have enlarged male eyes, characterized by a higher number of ommatidia compared with worker eyes and a frontal zone with enlarged facets, which is likely to be associated with improved spatial resolution and contrast sensitivity. In these species, perching mating strategies are found. In contrast, males of species that patrol scent routes (Bombus hortorum, Bombus lapidarius, Bombus pascuorum, Bombus pratorum, Bombus soroeensis, Bombus terrestris, Bombus wurflenii) show no distinct eye adaptations, and their eye morphology closely resembles that of the workers. A phylogenetic analysis of male eye structure indicates that enlarged males eyes may have been the ancestral state in Bombus. © 2014 The Linnean Society of London

Roving females and patient males: a new perspective on the mating strategies of chimpanzees

Mating strategies are sets of decisions aimed at maximizing reproductive success. For male animals, the fundamental problem that these strategies address is attaining mating access to females in a manner that maximizes their chances of achieving paternity. For chimpanzees (Pan troglodytes), despite substantial interest in mating strategies, very little attention has been paid to the most fundamental problem that mating strategies need to solve: finding mates. Only a single model, Dunbar's general model of male mating strategies, exists to explain mate-searching behaviour in chimpanzees. Under this model, males in most populations are regarded as pursuing a 'roving' strategy: searching for and sequestering fertile females who are essentially passive with respect to mate searching. The roving mating strategy is an assumption deeply embedded in the way chimpanzee behaviour is considered; it is implicit in the conventional model for chimpanzee social structure, which posits that male ranging functions both to monitor female reproductive state and to ward these females from other groups of males through collective territoriality: essentially, ranging as mating effort. This perspective is, however, increasingly at odds with observations of chimpanzee behaviour. Herein, I review the logic and evidence for the roving-male mating strategy and propose a novel alternative, a theoretical framework in which roving is a strategy pursued by female chimpanzees in order to engage successfully in promiscuous mating. Males, unable to thwart this female strategy, instead maximise the number of reproductive opportunities encountered by focusing their behaviour on countering threats to health, fertility and reproductive career. Their prolonged grooming bouts are seen, in consequence, as functioning to mitigate the negative impacts of socially induced physiological stress. In this new framework, the roving-male strategy becomes, at best, a 'best of a bad job' alternative for low-ranking males when faced with high levels of competition for mating access. Male chimpanzees do not search for mates, but for one another, for food, and, at times, for rivals in other communities. To the extent that female promiscuity functions to counter infanticide risk, mate searching by female chimpanzees-and any associated costs-can be seen as an unavoidable consequence of male sexual coercion. This novel framework is a better fit to the available data than is the conventional account. This review highlights the desperate need for additional work in an area of chimpanzee biology that has been somewhat neglected, perhaps in part because assumptions of roving males have remained unquestioned for too long. It also highlights the need, across taxa, to revisit and revise theory, and to test old assumptions, when faced with contrary data.

Trade-off between resource allocation and acquisition in anadromous adult male Atlantic salmon (Salmo salar L.)

Trade-off between resource allocation and acquisition in anadromous adult male Atlantic salmon (Salmo salar L.)

A dominant allele controls development into female mimic male and diminutive female ruffs

Maintaining polymorphisms for genes with effects of ecological significance may involve conflicting selection in males and females. We present data from a captive population of ruffs (Philomachus pugnax) showing that a dominant allele controls development into both small, ‘female mimic’ males (‘faeders’), and a previously undescribed class of small ‘female faeders’. Most male ruffs have elaborate breeding plumage and display behaviour, but 0.5–1.5% are faeders, which lack both. Females from a captive population previously lacking faeders were bred with two founder faeder males and their faeder sons. The faeders’ offspring had a quadrimodal size distribution comprising normal-sized males and females, faeders and atypically small females. By contrast, ornamented males fathered only normal-sized offspring. We conclude that both founding faeders were heterozygous for a faeder allele absent from the original population. This allele is dominant to previously described genes that determine development into independent versus satellite ornamented males. Unlike those genes, the faeder allele is clearly expressed in females. Small body size is a component of the male faeder mating strategy, but provides no obvious benefit to females. Bisexual expression of the gene provides the opportunity to quantify the strength of sexually antagonistic selection on a Mendelian trait.

Large donors transfer more sperm, but depletion is faster in a promiscuous hermaphrodite

Males are limited in sperm and seminal fluid, which can lead to prudent sperm allocation across consecutive matings. While sperm depletion is usually considered a population characteristic, we investigated an individual size-related variation in sperm depletion in the promiscuous sea slug Chelidonura sandrana. We found that sperm counts substantially declined across four successive copulations, indicating sperm depletion. Sperm depletion occurred at a disproportionally fast rate in larger sperm donors. Based on our finding of body size-dependent sperm depletion, we further investigated whether this translates into prudent sperm allocation across ejaculates. We found no evidence that slugs adjusted the number of delivered sperm in response to partner body weight, a tight proxy of fecundity. Instead, sperm counts increased with sperm donor body weight. Sperm donation independent of partner body size and—as found in previous work—partner novelty leaves scope for currently unidentified traits like the sperm competition environment to affect male sperm allocation. Alternatively, we propose that, in C. sandrana, a low optimal male mating rate close to the female optimum as indicated by our data may ultimately render indiscriminate sperm allocation the most beneficial male mating strategy.

A unifying framework reveals key properties of multilevel selection

Natural selection can act at multiple biological levels, often in opposing directions. Viral evolution is an important example, with selection occurring both within infected hosts and between hosts via transmission. A fast-replicating virus may outcompete a slower strain within the same host, however, if rapid viral replication incapacitates the host, this fast-replicating virus may not be transmitted as frequently as its slower counterpart. Such examples of antagonistic multilevel selection arise across biological taxa and scales, from microbial public goods production to male mating strategies. A general formalism for describing and analyzing these diverse systems can identify their common underlying properties. Here I introduce such a unifying framework, which can be intuitively visualized as a stochastic ball-and-urn process. This ball-and-urn process illustrates the dynamics of antagonistic selective forces and allows the systematic derivation of properties with little or no dependence on model parameterization. These properties are consistent with previous studies, both theoretical and empirical, of multilevel selection. In particular I show that selection at the group level is favored when group-level events occur frequently relative to individual-level events, when there is little or no mutation, and when there are many groups relative to the number of individuals in each group. This approach demonstrates how multilevel selection can be understood as a general biological phenomenon, and identifies recurring characteristics that may be independent of specific biological contexts.

The Dark Triad: Beyond a ‘male’ mating strategy

The Dark Triad (DT: sub-clinical narcissism, Machiavellianism and psychopathy) is argued to facilitate a male short-term mating strategy. The trait constellation in women and its potential adaptive benefits has received less attention. We examined the prevalence and correlates of DT in a large community sample (N=899). Despite finding expected sex differences in Sensation-seeking, Competitiveness, strength of sexual motivation, recreational sex behaviors and neuroticism, we found no sex difference in DT scores. Furthermore, within-sex multiple regressions identified the same predictors of DT score with similar weightings. Moderation analysis confirmed regression equations did not differ by sex. We propose that focus on DT as a male adaptation to short-term mating has been overstated and that men’s greater preference for casual sexual encounters is not explained by DT traits.

Why Male Mammals Are Monogamous

Social monogamy evolved many times across mammals as a male mating strategy in species where females are widely spaced for ecological reasons. [Also see Research Article by Lukas and Clutton-Brock ]

Intersexual dominance relationships and the influence of leverage on the outcome of conflicts in wild bonobos (Pan paniscus)

Dominance relationships between females and males are characteristic traits of species and are usually associated with sexual dimorphism. Exploring the social and contextual circumstances in which females win conflicts against males allows one to study the conditions triggering shifting power asymmetries between the sexes. This study investigates dominance relationships in bonobos (Pan paniscus), a species in which females are thought to display social dominance despite male-biased sexual dimorphism. To identify dominance relationships among females and males, we first explored how intrasexual dominance status affects the outcome of intersexual conflicts. Second, by incorporating social and behavioral information about the context of intersexual conflicts, we tested to which extent different components of power are relevant to the observed asymmetries in the relationships. Post-hoc analyses indicate a sex-independent dominance hierarchy with several females occupying the top ranks. Our results also reveal that two factors—female leverage and motivation to help offspring—had a significant influence on the outcome of intersexual conflicts. The results of our study do not indicate an overall reduction in male aggression against females but do show lower levels of male aggression in the mating context, and an absence of male aggression toward those females displaying visual signs of elevated fecundity. This indicates that both female sexuality and male mating strategies are involved in the shifting dominance relationships between the sexes.

Male coalitions and female behaviour affect male mating success independent of dominance rank and female receptive synchrony in wild Barbary macaques

Dominant mammalian males should gain a reproductive advantage due to their greater fighting abilities. However, the extent to which they can monopolise access to females varies across species. In primates and recently other mammalian species, the Priority of Access (PoA) model is commonly used to measure the degree to which male rank and female receptive synchrony affect mating skew. Few studies have examined the factors which lead to deviations from the expectations of the model. Here, we investigate male mating skew in wild Barbary macaques (Macaca sylvanus). We examined four of the main factors which affect male mating success: the roles of male rank, female receptive synchrony, coalitionary activity and female behaviour. We found that male mating was skewed up the hierarchy, but there was a large deviation from the PoA model's expectations with high-ranked males not gaining as big a share as expected. Females frequently initiated sexual encounters, predominantly with mid-ranked males, increasing their mating success. Male coalitionary activity independently increased mating success. Frequent associations with females were costly to males as they were the targets of bridging coalitions, decreasing future mating opportunities for the targets. High-ranking males did not increase their mating success directly through bridging coalitions but acted to dilute the effects of female behaviour. By examining different factors affecting mating skew, we are able to show that alternative male and female mating strategies are effective in reducing the monopolisation potential of the dominant male.

Consequences of snowy winters on male mating strategies and reproduction in a mountain ungulate

Alternative mating tactics (AMTs) are intrasexual variants in mating behaviour of several species ranging from arthropods to mammals. Male AMTs coexist between and within populations. In particular, male ungulates rarely adopt just one tactic throughout their lifetime. Tactics commonly change according to internal factors (age, body size, condition) and external conditions (weather, resources, predation, animal density). However, the influence of weather has not yet been investigated in upper vertebrates. Such influence may be relevant in species whose rutting period occurs late in fall or in winter, when environmental conditions and the snow cover in particular may vary considerably. We detected two AMTs in Alpine ibex (Capra ibex) males: older and full-grown males mainly adopted the tending tactic, while younger males usually pursued an alternative one (coursing tactic). Weather was found to influence the use of AMTs by males: in snowy mating seasons, the coursing tactic was no longer used due to difficulties in moving through deep snow. In snowy rutting periods, males appeared to delay or even avoid mating activities and a decrease of births was reported in the second part of the following birth season. Snow cover may have a negative effect on population dynamics by reducing the recruitment and on population genetic variability, as a consequence of poorer mating opportunities. Studies on factors affecting mating behaviour and leading to a reduced availability of mates and a decrease in female productivity are especially relevant in species, like Alpine ibex, whose genetic variability is low.

Experimental tests of mate choice in nonhuman mammals: the need for an integrative approach

Experimental studies of mate choice have normally focused on non-mammal animal species, in which female mating preferences are based on clearly defined male traits. Because mammals are invariably larger and behaviourally more complex, they are less suited to this type of experimentation. Nevertheless, numerous studies on nonhuman mammals have shown that females appear to actively choose their mates. In this Commentary, I review the current literature to reveal that most experimental tests of mate choice in mammals are unable to reveal the actual male phenotypic trait(s) of female preference, which is crucial for identifying male characteristics under sexual selection. In addition, very few studies take into account female oestrous stage, or quantify the fitness benefits to discriminating females. Future work should concentrate on demonstrating female preferences for specific male traits that are shown by genetic paternity analysis to be correlated with male reproductive success, using setups that control for the effects of male and female mating strategies and in which the actual experiments are performed during the female's peak oestrous period.

Mating behavior and birth seasonality of black-and-white snub-nosed monkeys (<I>Rhinopithecus bieti</I>) at Mt. Lasha

Copulation patterns are important to understanding male mating strategies and stabilization strategies of social organizations in primates. However, information on copulation patterns of Asian primates is relatively rare. This study was undertaken to collect data on mating behavior and birth seasonality of Black-and-white Snub-nosed monkeys (Rhinopithecus bieti) using all occurrence sampling and Focal animal-scan sampling methods at Mt. Lasha, between January and December, 2011. Our study focused on observing mating frequency and birth rates. Snub-nosed monkeys mate year round, with two observable peaks: one reproductive peak during the mating season, roughly from August to October, and a second non-breeding peak during the birth season. It is unclear what biological significance this non-reproductive mating peak has. During our observation, we noted a lower ratio of mount to ejaculation and rare ejaculatory copulations, which indicated that every mating would not result in ejaculation. This study corroborates the previous view that the Rhinopithecus bieti's copulatory pattern is likely multiple-mount ejaculation (MME) or at the upper part of mating continuum of single-mount ejaculation (SME) toward MME. More ejaculatory copulations initiated by males than females indicate that MME is not only a mating strategy of males, but that males can influence the position of their copulatory pattern on the continuum between SME and MME. The mating frequencies significantly correlated with the birth rates with a delay of 6 to 7 months. Monkeys gave birth within a strict seasonality with a birth peak of March, which confirms the previous view. Infants were born with a certain degree of synchronization, but different populations displayed different modes of synchronizations.

An ecological analysis of mating biology of Xylocopa virginica in southern Ontario

1. Xylocopa virginica virginica Linnaeus is a wide‐ranging species with plastic nesting behaviour that appears to represent an intermediary between solitary and social nesting species. Over 3 years, a natural population was studied with the objective of quantifying the relationship among population dynamics, climate, female nest provisioning behaviour, and male mating strategy.2. Males in the population congregated around female‐occupied nesting sites before the beginning of nest provisioning by females; both resident and satellite male mating strategies were observed. Overall, the present results are consistent with female defence polygyny.3. Male mating strategies were consistent across the three breeding seasons of our study, in spite of annual variation in population size, sex ratio, and weather. Male mating behaviour was also consistent with that seen in other populations with longer breeding seasons.4. Adult non‐breeding females that never leave nests are observed in nests throughout the breeding season and we hypothesise that males continue to defend territories after breeding females have mated because of a small probability they can mate with one of these non‐breeding females.5. These results are important to our understanding of the relationship between mating systems and the evolution of sociality, contributing data on the role of ecological factors to male mating behaviour. Collection of such data for a variety of species that differ in sociality is necessary for the comparative analysis that is required to fully elucidate coevolution of mating systems and sociality.

Influence of density and major histocompatibility genotype on sexual selection in a salmonid alternative mating strategy

Various mechanisms of sexual selection can occur at both the whole organism and gamete levels. Fertilization success in salmonid fishes is largely determined by behavioural competition within and between “fighter” and “sneaker” male strategies, but is also influenced by interactions among gametes. We investigated the influence of density, fighter male presence, and major histocompatibility (MH) genotype on patterns of fertilization success in sneaker Atlantic salmon ( Salmo salar ). At low density and in the absence of fighter males, monopolization by single sneaker males occurred, suggesting that male–male competition was the main determinant of fertilization success. By contrast, at high density and (or) in the presence of fighter males, several sneakers succeeded in fertilizing eggs. Our study, the first to investigate the role of MH-mediated fertilization for a genetically based alternative male mating strategy, suggests that a larger proportion of eggs were fertilized by sneakers whose MH genotypes were more similar to those of the female than expected by chance. These findings highlight the importance of examining sexual selection under different competitive conditions.

Under the Hood of Tess: Conflicting Reproductive Strategies in Thomas Hardy’s Tess of the D’Urbervilles

Thomas Hardy’s Tess of the D’Urbervilles is analyzed from an evocritical perspective in order to consider evolved human reproductive strategies through the psychology and behavior of the novel’s three principal characters: Tess, Alec and Angel. It is argued that Hardy made the episode of Tess’ and Alec’s sexual contact, as well its interpretation by the characters, ambiguous, thereby suggesting the possibility of seduction rather than rape. In this context, two female mating patterns—inherited from our hominid ancestors—appear in Tess’ behavior: (a) the collection of high quality genes from a genetically fit male (Alec) who is not likely to stay with the female and provide for the offspring and (b) mating with a provider male who is interested in long-term parental investment (Angel). Conversely, Angel and Alec represent two male mating strategies that evolved as possible courses of action in our species: the dad and the cad respectively. The unwillingness of Angel to forgive Tess her sexual past is considered in the context of another evolved feature of the human mind: paternal uncertainty (the fear of the male’s genetic extinction through the possibility of raising another male’s offspring). This is juxtaposed with studies of male jealousy in different cultures and periods. Tess’ decision to tell Angel about her past is viewed in connection with the concept of modularity: an approach to human psychology based on the assumption that the mind is divided into specialized modules (responsible for different cognitive spheres) which can sometimes conflict.

Complex patterns of male alliance formation in a dolphin social network

The formation and maintenance of alliances is regarded as one of the most socially complex male mating strategies in mammals. The prevalence and complexity of these cooperative relationships, however, varies considerably among species as well as within and between populations living in different ecological and social environments. We assessed patterns of alliance formation for Indo-Pacific bottlenose dolphins, Tursiops aduncus, in Port Stephens, New South Wales, Australia, to investigate the stability of these alliances, the structure of associations, as well as variation in schooling patterns among males. Our results showed that association patterns among males within this population showed considerable variability. Males either formed strong and enduring alliances that lasted for at least 8 years with minimal partner switching, or less stable partnerships within a much larger male social network. Male alliances with the strongest levels of association within a given time period were significantly more likely to maintain their relationships over the long term compared with alliances with lower levels of association. Males in stable alliances also associated in significantly smaller schools than males who formed less stable alliance partnerships. Finally, we found that alliances consisting of more related males did not persist longer than alliances between unrelated individuals. Our study suggests that intrapopulation variation in male alliance formation in dolphins likely reflects different mating strategies adopted as individual responses to their complex fission–fusion social environment.

Determinants of Natural Mating Success in the Cannibalistic Orb-Web Spider Argiope bruennichi

Monogynous mating systems (low male mating rates) occur in various taxa and have evolved several times independently in spiders. Monogyny is associated with remarkable male mating strategies and predicted to evolve under a male-biased sex ratio. While male reproductive strategies are well documented and male mating rates are easy to quantify, especially in sexually cannibalistic species, female reproductive strategies, the optimal female mating rate, and the factors that affect the evolution of female mating rates are still unclear. In this study, we examined natural female mating rates and tested the assumption of a male-biased sex ratio and female polyandry in a natural population of Argiope bruennichi in which we controlled female mating status prior to observations. We predicted variation in female mating frequencies as a result of spatial and temporal heterogeneity in the distribution of mature females and males. Females had a low average mating rate of 1.3 and the majority copulated only once. Polyandry did not entirely result from a male-biased sex-ratio but closely matched the rate of male bigamy. Male activity and the probability of polyandry correlated with factors affecting pheromone presence such as virgin females' density. We conclude that a strong sex ratio bias and high female mating rates are not necessary components of monogynous mating systems as long as males protect their paternity effectively and certain frequencies of bigyny stabilise the mating system.