Male aggression during mating: evidence for sexual coercion in a female dominant primate?

In this article we document male sexual coercion of a Lemur catta female on St. Catherines Island (SCI), USA. Data presented in this paper were collected on one free-ranging L. catta group during October-November 2002 using all-occurrences sampling for agonism and reproductive behavior. We observed a male forcefully attempting to mate with a year-old estrous female. Despite the fact that we observed this female to present to the male during her estrus, throughout the episode the male employed the use of force, and achieved penile intromission on at least one occasion while the female struggled and resisted. We interpret his behavior as sexual coercion. As measured by sexual presents, the female appeared to more strongly prefer two other males as mates, yet each of these males gained fewer mounts and less cumulative time spent in mounts than the coercive male. The coercive male was one of two group males with the lowest observed mating success, suggesting that coercion might be a strategy used by males who are not highly sexually preferred. Females undergo early sexual maturation at this site owing to provisioning, and can therefore enter estrus before fully attaining dominance over males. As such, these data suggest that one consequence of provisioning a wild L. catta population (or of maintaining L. catta in captivity) may be that young females can be the targets of sexual coercion if they reach sexual maturity before fully achieving social dominance over males. In conclusion, male sexual coercion can occur in L. catta despite the female dominance characteristic of this species, and can constrain the mating behavior of females.     

Sexual harassment in live-bearing fishes (Poeciliidae): comparing courting and noncourting species

Sexual harassment by males has been reported from several live-bearingfishes (Poeciliidae) and has been shown to inflict costs onfemales. For example, poeciliid females have reduced feedingopportunities when accompanied by a male because females dedicateattention to avoiding male copulation attempts. Poeciliid speciesdiffer considerably in male mating behavior, such as the presenceor absence of courtship. Courting males display in front ofthe females, but males attempting to sneak-copulate approachfemales from behind, that is, in the blind portion of theirvisual field, and force copulations, which can be viewed asa male persistence trait. We predicted that poeciliid femalesneed to be more vigilant in the presence of noncourting males,and costs of harassment by noncourting males might be stronger.In a comparative approach we examined the costs of male sexualharassment for females as reduced feeding time in 9 speciesof live-bearing fishes, including courting (Poecilia latipinna,Poecilia reticulata, Xiphophorus cortezi, Xiphophorus variatus)and noncourting species (Poecilia mexicana [surface- and cave-dwellingform], Poecilia orri, Gambusia affinis, Gambusia geiseri, Heterandriaformosa). In all species examined except for the cave form ofP. mexicana, focal females spent significantly less time feedingin the presence of a male than when together with another female.The time females spent feeding was found to significantly declinewith increasing male mating activity (sum of all sexual behaviors),but there was no support for the idea that females would spendmore time feeding in the presence of courting males comparedwith noncourting ones.     

Sexually antagonistic coevolution in insects is associated with only limited morphological diversity

Morphological traits involved in male-female sexual interactions, such as male genitalia, often show rapid divergent evolution. This widespread evolutionary pattern could result from sustained sexually antagonistic coevolution, or from other types of selection such as female choice or selection for species isolation. I reviewed the extensive but under-utilized taxonomic literature on a selected subset of insects, in which male-female conflict has apparently resulted in antagonistic coevolution in males and females. I checked the sexual morphology of groups comprising 500-1000 species in six orders for three evolutionary trends predicted by the sexually antagonistic coevolution hypothesis: males with species-specific differences and elaborate morphology in structures that grasp or perforate females in sexual contexts; corresponding female structures with apparently coevolved species-specific morphology; and potentially defensive designs of female morphology. The expectation was that the predictions were especially likely to be fulfilled in these groups. A largely qualitative overview revealed several surprising patterns: sexually antagonistic coevolution is associated with frequent, relatively weak species-specific differences in males, but male designs are usually relatively simple and conservative (in contrast to the diverse and elaborate designs common in male structures specialized to contact and hold females in other species, and also in weapons such as horns and pincers used in intra-specific battles); coevolutionary divergence of females is not common; and defensive female divergence is very uncommon. No cases were found of female defensive devices that can be facultatively deployed. Coevolutionary morphological races may have occurred between males and females of some bugs with traumatic insemination, but apparently as a result of female attempts to control fertilization, rather than to reduce the physical damage and infections resulting from insertion of the male's hypodermic genitalia. In sum, the sexually antagonistic coevolution that probably occurs in these groups has generally not resulted in rapid, sustained evolutionary divergence in male and female external sexual morphology. Several limitations of this study, and directions for further analyses are discussed.     

Mate sampling and the sexual conflict over mating in seaweed flies

The order in which females encounter, or sample, males in apopulation may have important consequences for mate choice,with the information gathered about males influencing boththe preference function and degree of choosiness of females.Sexual selection may be affected as a result. Sampling of particularsubsets of males may be a crucial component of individual variation in mate preferences within populations. However, the sequencein which males are sampled may also be important in specieswithout traditional, active mate choice, such as when sexualselection involves sexual conflict over mating. This wouldoccur if the likelihood of a female mating with a male of acertain phenotype changes as a result of previous encounters.We examined the effects of encountering males differing inbody size, a sexually selected phenotype, in the seaweed flyCoelopa frigida. Sexual selection occurs in this species asa result of a sexual conflict over mating. We show that theoutcome of the sexual conflict is independent of the orderin which males are encountered by female seaweed flies, withthe overall mating advantage to large males being unaffected.In addition, we explored female preference functions and evaluatethe heterogeneity in female willingness to mate. We suggestthat consideration of mate sampling theory is valuable whenexamining mate choice in species in which sexual selectionis driven by sexual conflict.     

Sexual dimorphism and dichromatism in Steere's Liocichla (Liocichla steerii)

ABSTRACT.   Although sexual differences in birds can be extreme, differences between males and females in body size and plumage color are more subtle in many species. We used a genetic-based approach to determine the sex of male and female Steere's Liocichla ( Liocichla steerii ) and examine the degree of size dimorphism and plumage dichromatism in this apparently monomorphic species. We found that males were significantly larger than females. In addition, Steere's Liocichla have a prominent yellow plumage patch on the lores that was significantly larger in males than females for both live birds and museum specimens. We also used reflectance spectrometry to quantify the color of the yellow-green breast feathers of Steere's Liocichla and found no significant differences between males and females in brightness, intensity, saturation, or hue. However, females tended to have brighter breast plumage, particularly at long wavelengths. Collectively, these color variables were useful in discriminating birds according to sex when used in a discriminant function analysis. Our study suggests that sexual selection may be more widespread than once assumed, even among birds considered monomorphic, and emphasizes the need for additional data from tropical and subtropical species.     

Conflict resolution in the Odonata: implications for understanding female mating patterns and female choice

Predictions of mating patterns in animals have focused on males and how they compete for fertilizations by controlling females. With reference to the Odonata, a taxon in which mating requires cooperation of the female, the active role that females play in mating decisions is often ignored, leading to the premature conclusion that male coercion of females is common. A critical review of the outcome of sexual conflict among odonates leads me to alternative explanations of female mating patterns that need to be refuted before concluding that males coerce matings. Because Anisoptera males have greater control over tandem formation, they have a greater potential for coercion than Zygoptera males. However, Anisoptera females may simply be willing to remate more often if they receive insufficient sperm to fertilize an entire egg clutch. Contrary to prior assumptions, in both suborders, male defence of oviposition sites does not preclude females from choosing among sites or among males. I find that the evolution of non-aggressive sexual signals by males is a reliable indication that sexual conflict has been resolved in favour of female interests. Although I predict that the benefits to females of choice of male phenotype should rarely exceed the cost of such discrimination in Odonata, female choice is most likely to evolve in territorial species whose males must endure high physiological stress in order to mate, and when site quality is not a reliable predictor of the genetic quality of a potential mate.     

Allometry and sexual selection of male weaponry in Wellington tree weta, Hemideina crassidens

Both male and female Wellington tree weta, Hemideina crassidens,use cavities in trees as diurnal shelters. That these galleriesare often limiting in nature offers males the opportunity toincrease their reproductive success by monopolizing galleriesand the females residing in them. Male H. crassidens, can matureat either the 8th, 9th, or 10th instar, whereas females matureat the 10th instar only, and male head (and mandible) size positivelycovaries with ultimate instar number. It has been suggestedthat males fight for control of galleries by using their enlargedmandibles as weapons, and males with larger mandibles controlgalleries with more females. In the present study, I presenta statistical examination of sexual dimorphism, showing thattraits related to head size are on average significantly largerin males, whereas traits related to body size are on averagesignificantly larger in females. I tested three predictionsaddressing the hypothesis that sexual selection is driving megacephalyin male H. crassidens. First, as predicted, traits related tohead size show a positive allometric relationship with bodysize in males but not in females. Second, adapting a novel statisticaltechnique based on maximum likelihood and bootstrapping revealedthat males, but not females, exhibit a multimodal distributionin head and body size traits. This is likely a consequence ofmales maturing at one of three instars, which results in positivecovariance between the ultimate instar number and morphologicaltraits. Third, as predicted, single adult males with largerheads reside in galleries housing larger groups of adult females.     

Mate choice based on static versus dynamic secondary sexual traits in the dark-eyed junco

Some secondary sexual traits (SSTs) such as structural characteristicsare semi-permanent or static, while others, such as courtshipdisplay, are more labile or dynamic. In this paper we reportresults from two experiments designed to test the relative attractivenessto female dark-eyed juncos (Junco hyemalis, Passeriformes, Aves)of a relatively static plumage trait, the amount of white inthe tail, and a relatively dynamic behavioral trait, courtshipintensity. The experiments derived from a study showing that femalejuncos prefer males that court more vigorously. We asked whether femalesalso base their preferences on plumage traits and how they respond whenpresented with a choice between attractive traits that are eitherstatic (plumage) or dynamic (courtship) in nature. In the firstexperiment we presented males to females in paired mate-choicetrials and found that males enhanced with more white in theirtails were more attractive to females than controls with unenhancedtails. Females spent more time with enhanced males and directedmore sexual displays toward them. In the second experiment we testedwhether females preferred males with enhanced tails (a staticSST) or males with enhanced hormone-mediated courtship behavior(a dynamic SST). In this experiment females did not demonstratea consensus preference for either the static or the dynamictrait. Instead, some females preferred the male whose courtshipperformance was enhanced with testosterone, while others preferredthe male with an enhanced tail. We conclude that both kindsof traits are important in junco mate choice, but that somefemales apparently weigh static traits more heavily than dynamicones, while other females use opposite weightings.     

Nuptial gift in the spider Pisaura mirabilis maintained by sexual selection

The nuptial prey gift in the spider Pisaura mirabilis has been suggested to function as a male protection against sexual cannibalismduring courtship and mating. This hypothesis together withtwo alternatives—male mating effort and paternal investment hypotheses—were tested in a laboratory experiment withsexually inexperienced males and females. One group of malesoffered no gift to the female while three groups of males offeredsmall, medium, or large sized gifts, respectively. No malewas cannibalized among 82 trials. Aggression was observed onlyin encounters where a gift was presented. Males without a gift courted females, and 40% of these males managed to copulate,compared to 90% of males offering a gift. The copulation durationwas positively correlated with gift size. In general, the femaleterminated the copulation and ran away with the gift. The proportionof eggs fertilized increased with copulation time. Presenceor size of the nuptial gift did not affect female fecundityor spiderling size significantly. The results refute the hypothesesof sexual cannibalism and paternal investment. The nuptialgift represents a male mating effort; it entices the femaleto copulate, facilitates coupling during copulation, and byprolonging copulation it may increase the amount of sperm transferred.I conclude that the nuptial prey gift in Pisaura mirabilisis maintained by sexual selection.     

Sexual Competition,Coercive Mating and Mate Assessment in the One‐Sided Livebearer,Jenynsia multidentata: Are They Predictive of Sexual Dimorphism?

We investigated the mechanisms of sexual selection operating on body size in the one‐sided livebearer (Jenynsia multidentata), a small fish characterized by male dwarfism. Mating in the one‐sided livebearer is coercive: males approach females from behind and try to thrust their copulatory organ at the female genital pore. Females counter males' mating attempts by either swimming away or attacking them. We tested the hypothesis that the components of sexual selection favouring small size in males (sexual coercion) were more effective than those favouring a large size (male competition and mate choice). When alone, small males had a significantly higher success in their mating attempts than large males. The proportion of successful attempts was also positively correlated with female size. When two males competed for the same female, the large male had a significant mating advantage over the small one. With a 1 : 1 sex ratio, the large‐male mating advantage vanished because each male tended to follow a different female. Large males, however, preferentially defended large females, thus compelling small males to engage with smaller, less fecund females. Males did not discriminate between gravid and non‐gravid females, but preferred mating with larger females. This preference disappeared when males were much smaller than the female, probably in relation to the risk for the male of being eaten or injured by the female. In a choice chamber, male‐deprived females that had their sperm storage depleted remained close to males and showed a preference for large individuals, a behaviour not observed in non‐deprived females. Nonetheless, when placed with males in the same aquarium, all females showed avoidance and aggression. Struggling may represent a way by which the female assesses the skill and endurance of males.     

Does courtship behavior contribute to species-level reproductive isolation in field crickets?

Reproductive behavior influences gene flow within and amongspecies; thus, sexual selection may be a major contributor tothe maintenance of species, and possibly their formation. HereI experimentally manipulate the courtship interactions of thefield crickets Gryllus rubens and G. texensis to examine thepotential of close-range courtship interactions to limit interspecificgene flow. Responses of males to females and of females to malecourtship song and males per se were examined for four pairedsympatric and allopatric populations. Male G. rubens were morelikely to court conspecific females, but male G. texensis courtedfemales of both species equally. If paired with conspecificmales, female G. rubens and G. texensis both preferred conspecificcourtship song. In none of these comparisons were the responsesof males or females from allopatry different from those in sympatry.There was an asymmetry of courtship response across both sexand species: male G. rubens were more discriminating than maleG. texensis, whereas female G. texensis were more discriminatingthan female G. rubens. Despite significant preferences for conspecifics,the net effect of courtship interactions would appear to limitinterspecific gene flow only weakly. These results are consistentwith courtship behavior evolving under the influence of sexualselection and only indirectly promoting species integrity.     

Sex and Weapons: Contrasting Sexual Dimorphisms in Weaponry and Aggression in Snapping Shrimp

Sexual dimorphisms in weaponry and aggression are common in species in which one sex (usually males) competes for access to mates or resources necessary for reproduction – sexually dimorphic weaponry and aggression, in other words, are frequently the result of intrasexual selection. In snapping shrimp, the major chela (snapping claw) can be a deadly weapon, and males of many species have larger chelae than females, a pattern readily interpreted as resulting from intrasexual selection. Thus, males might be expected to show more sex‐specific aggression than females, and be more aggressive overall. We tested these predictions in two species of snapping shrimp in a territorial defense context. Neither of these predictions was supported: in both species, females, but not males, engaged in sex‐specific aggression and females were more aggressive than males overall. These contrasting sexual dimorphisms – larger weaponry in males but higher aggression in females – highlight the importance of considering the function of weaponry and aggression in contexts other than direct competitions over mates. In addition, species differences in the degree of sexual dimorphism in chela size were due to differences in female, not male, chela size, and the species with greater sexual dimorphism in weaponry was significantly less aggressive overall; also, while paired and solitary males did not differ in residual chela size, for the species with greater sexual dimorphism, females carrying embryos had smaller residual chela sizes. These results suggest that understanding the sexual dimorphisms in weaponry and aggression in snapping shrimp requires understanding the relative costs and benefits of both in females as well as males.     

Sexual cannibalism, competition, and size dimorphism in the orb-weaving spider Nephila plumipes Latreille (Araneae: Araneoidea)

The degree and direction of sexual dimorphism varies widely,but in several taxa of orb-weaving spiders, including Nephila,males may be less than one-tenth the size of females. This differenceis commonly attributed to selection through precopulation sexualcannibalism: females may either fail to detect very small males,or ignore them as potential prey items. However, there is oftenthe potential for male-male competition in these species becauseseveral males can be found on the web of a single female. Weinvestigated experimentally the effects of sexual cannibalismand male-male competition on male body size and hence sexualdimorphism in the Australian golden orb-weaver (Nephila plumipes).Small males were less likely to be detected and cannibalizedthan larger males. However, larger males excluded small malesfrom the central hub of the web, where mating takes place. Theconflicting effects of sexual cannibalism and male-male competitionmay be responsible for the relatively large variation in malebody size in this species.     

Safer sex with feeding females: sexual conflict in a cannibalistic spider

Mating strategies are to a large degree shaped by conflictsbetween the sexes, causing a rapid antagonistic coevolutionof traits involved in reproduction. The view that sexual cannibalismrepresents a form of sexual conflict leads to the predictionof male traits that facilitate escape from cannibalistic females.A variety of traits have been suggested to serve this functionin spiders, where sexual cannibalism is comparatively common.Empirical evidence, however, is virtually absent. Here we showexperimentally that opportunistic mating with feeding females,which has been reported from several species of orb-weavingspiders, greatly reduces the risk of cannibalism and injuryfor males in the spider Nephila fenestrata. This has directconsequences for a male's fertilization success because survivingmales can reduce the female's remating probability by guardingher against rivals. Although copulation with previously matedfemales sometimes appears to be mechanically impossible, secondmales that do copulate can expect to fertilize on average 64%of a female's eggs. Our results support the view that opportunisticmating may have evolved as a male tactic in a context of sexualconflict over sexual cannibalism.     

Female resistance to sexual coercion can evolve to preserve the indirect benefits of mate choice

Sexual conflict over the indirect benefits of mate choice may arise when traits in one sex limit the ability of the other sex to freely choose mates but when these coercive traits are not necessarily directly harmful (i.e. forced fertilization per se). Although we might hypothesize that females can evolve resistance in order to retain the indirect, genetic benefits (reflected in offspring attractiveness) of mating with attractive males, up to now it has been difficult to evaluate potential underlying mechanisms. Traditional theoretical approaches do not usually conceptually distinguish between female preference for male mating display and female resistance to forced fertilization, yet sexual conflict over indirect benefits implies the simultaneous action of all of these traits. Here, we present an integrative theoretical framework that draws together concepts from both sexual selection and sexual conflict traditions, allowing for the simultaneous coevolution of displays and preferences, and of coercion and resistance. We demonstrate that it is possible for resistance to coercion to evolve in the absence of direct costs of mating to preserve the indirect benefits of mate choice. We find that resistance traits that improve the efficacy of female mating preference can evolve as long as females are able to attain some indirect benefits of mating with attractive males, even when both attractive and unattractive males can coerce. These results reveal new evolutionary outcomes that were not predicted by prior theories of indirect benefits or sexual conflict.     

Development of intraspecific size variation in black coucals,white-browed coucals and ruffs from hatching to fledging

Most studies on sexual size dimorphism address proximate and functional questions related to adults, but sexual size dimorphism usually develops during ontogeny and developmental trajectories of sexual size dimorphism are poorly understood. We studied three bird species with variation in adult sexual size dimorphism: black coucals (females 69% heavier than males), white-browed coucals (females 13% heavier than males) and ruffs (males 70% heavier than females). Using a flexible Bayesian generalized additive model framework (GAMM), we examined when and how sexual size dimorphism developed in body mass, tarsus length and bill length from hatching until fledging. In ruffs, we additionally examined the development of intrasexual size variation among three morphs (Independents, Satellites and Faeders), which creates another level of variation in adult size of males and females. We found that 27–100% of the adult inter- and intrasexual size variation developed until fledging although none of the species completed growth during the observational period. In general, the larger sex/morph grew more quickly and reached its maximal absolute growth rate later than the smaller sex/morph. However, when the daily increase in body mass was modelled as a proportion, growth patterns were synchronized between and within sexes. Growth broadly followed sigmoidal asymptotic models, however only with the flexible GAMM approach, residual distributions were homogeneous over the entire observation periods. These results provide a platform for future studies to relate variation in growth to selective pressures and proximate mechanisms in these three species, and they highlight the advantage of using a flexible model approach for examining growth variation during ontogeny.     

Color patterns and species recognition in four closely related species of Lake Malawi cichlid

We examined interspecific female mating preferences in fourclosely related species of cichlid belonging to the Pseudotropheuszebra species complex of Lake Malawi. These species differin coloration but are similar in other respects, suggestingthat male color patterns may be important to female mate choicein species recognition. To test this hypothesis, we presented females from each species with a choice of four males, one ofher own species and three others that were each of a differentspecies. We also gave each female a choice between the threeheterospecific males only. In all four species, females showeda significant preference for conspecific males in the four-waychoice and chose the male with the most similar color patternto the conspecific male in the three-way choice. These resultsare discussed with reference to the theory of sexual selectionon color patterns as a means of sympatric speciation in cichlids.     

Sexual History Affects Mating Behavior and Mate Choice in the Crayfish Orconectes limosus

Because mating entails both costs and potential benefits to both sexes, males and females should be under selection to make optimal choices from among available potential mates. For example, in some cases, individuals may benefit by using information on potential mates' previous sexual histories to make mate choices. In such cases, the form and direction of these benefits may vary both between the sexes and based on the sexual history of the choosing individuals themselves. We investigated the effects of recent previous sexual history on the mate choice and mating behavior of both males and females of the crayfish Orconectes limosus. In one experiment, we found that opposite‐sex dyads comprising crayfish that had both mated 7–8 d previously with other conspecifics were significantly less likely to mate than dyads in which at least one crayfish was unmated. In a second experiment, we found that, when presented with a choice of tethered (but free to move) opposite‐sex conspecifics, only virgin females discriminated between males based on sexual history, showing a preference for virgin males over recently mated males. Mated females, mated males, and virgin males showed no preferences based on the sexual histories of potential mates. We discuss the implications of these inferences in the context of what was previously known about mating behavior and potential sperm limitation in crustaceans and other taxa.     

The effects of copulation duration in the bruchid beetle Callosobruchus maculatus

Control over copulation duration is a potentially importantgenerator of sexual conflict that has received little empiricalattention. The copulatory behavior of the bruchid beetle Callosobruchusmaculatus may reflect a sexual conflict over copulation duration.Males have spines on their intromittent organs that puncturethe female reproductive tract, and females kick their matesduring copulation. If females are prevented from kicking, copulationslast longer and the injuries females sustain are more severe.Males supposedly use the spines as anchors to prolong copulationduration, and females kick to terminate copulations. We manipulatedcopulation duration experimentally and quantified its effectson male and female fitness components to test whether or notthere is a conflict over copulation duration in C. maculatus.Females did not suffer from long copulations but instead experiencedincreased lifetime fecundity. Ejaculate size increased withcopulation duration, and females apparently derive materialbenefits from the ejaculates. Males that mated first and hadlong copulations were relatively unsuccessful when competingwith sperm from other males. However, there was a trend forfemale remating propensity to decrease with long copulationdurations, and first males may therefore also benefit from longcopulations. The copulation duration of the second male to matedid not have a significant effect on sperm precedence. We concludethat even though it seems likely that the male spines have evolvedto act as an anchor during copulation, there seems to be littleconflict over copulation duration per se in C. maculatus.     

Sexual selection in a moth: effect of symmetry on male mating success in the wild

Sexual selection is generally caused by female choice and male–malecompetition. In female choice process, female preference isfavored indirectly and/or directly by sexual selection. In indirectselection, females expressing the preference might gain indirectgenetic benefits. In direct selection, females expressing thepreference might gain direct benefits or avoid male-imposedcosts. The white-tailed zygaenid moth Elcysma westwoodii ismonandrous, and males often gather around a female to mate withher, suggesting a high opportunity for sexual selection on maletraits. We quantified phenotypic selection on male morphologyin this species in the field. The morphological characters analyzedincluded body weight, antenna length, forewing length, hindwing length, hind wing tail length, genital clasper length,and the fluctuating asymmetry (FA) of these bilateral traits.In E. westwoodii, selection favored males with more symmetricgenital claspers, as well as longer and more symmetrical hindwings and antennae. Negative correlations between FA and sizewere also detected in the clasper and the antenna. Our resultssuggest that FAs of male traits, in particular the genital clasper,may have indirect and direct influences on mating success. Duringa copulatory attempt, an E. westwoodii male will try to graspthe female's abdominal tip with his claspers but often failto do so because of the female's reluctance to mate. The femaleabdominal tips are smooth and strongly sclerotized and couldthus be difficult for males to grasp. We hypothesize that moresymmetrical male claspers are more efficient in overcoming femalereluctance.