Social and hormonal influences on behavior of adult male, female, and pseudohermaphroditic rhesus monkeys

We previously demonstrated that in a simple pair test situation the expression of adult male sexual behavior by rhesus monkeys depends on both prenatal (organizational) and adult (activational) androgen exposure. In the present study we used a more complex social situation (trio tests) to evaluate the behavior of males, females, and female pseudohermaphrodites. In these trio tests, the experimental subjects were tested with two estrogenized stimulus females simultaneously. Sex differences in behavior were made apparent by this complex testing situation that could not have emerged in the pair test. Gonadectomized males and female pseudohermaphrodites, but not ovariectomized females that were concurrently receiving TP, exhibited increased male sexual behavior in trio tests compared to pair tests. In trio tests, the males and pseudohermaphrodites showed evidence of partner preference by interacting almost exclusively with one of the two stimulus females. These "preferred females" in turn were responsible for the majority of the proceptive behavior exhibited in these tests. Ovariectomized females rarely displayed male sexual behavior in either test situation. These results further support the hypothesis that prenatal androgen exposure predisposes monkeys to exhibit masculine behavior traits when they reach adulthood and are exposed to the activational influences of androgens.     

Female mating preference for own males on species and population level in Chromaphyosemion killifishes (Cyprinodontiformes, Nothobranchiidae)

Chromaphyosemion killifishes are a very promising taxon for the examination of general principles and mechanisms underlying speciation. The polygamous mating system and high degree of sexual dimorphism in this group suggest that sexual selection and female mating preferences play a crucial role in the current radiation process. Because the emergence of mating preferences for individuals of the own population during allopatry is a necessary precondition for speciation by sexual selection, either via diverging mate recognition traits or reinforcement, we examined whether females of different nominal species (Chromaphyosemion bivittatum vs. Chromaphyosemion volcanum) or populations (C. bivittatum “Toko” vs. C. bivittatum “Ilor”) prefer their own males as mating partners. For this purpose, two different methods were used. First, we conducted simultaneous choice tests where females could choose between two different males, and second, we counted the eggs produced by females in consecutive matings with different males. Both approaches produced the same result: females of the populations under examination preferred to mate with their own males. Preference for own males was symmetrical in all tests. Males, on the other hand, do not discriminate between females, not even on the level of nominal species. Thus, our results are in accordance with the hypothesis that female choice is an important factor for speciation in Chromaphyosemion. Additionally, we present some preliminary data on post-zygotic isolation between the populations.     

Behavioral effects of a synthetic mixture of aliphatic acids in rhesus monkeys (Macaca mulatta)

The stimulation of sexual behavior by a synthetic mixture of volatile aliphatic acids (acetic, propanoic, methylpropanoic, butanoic, methylbutanoic, methylpentanoic) was studied in male rhesus monkeys. Twelve intact adult males and 12 long-ovariectomized adult females were used in 24 paired combinations (541 tests each of 1 hr). A mixture of authentic acids similar to that found in the vaginal secretions of estrogenized females was applied to the sexual skin area of ovariectomized females immediately before tests with males. There was marked between-pair variability during the application of both control and test substances. However, using rigorous behavioral criteria, there was a well-marked stimulation either of male mounting attempts or of ejaculations in 12 of 24 pairs involving 9 of 12 males. Three males responded with both female partners, three responded with neither female partner, and six responded with one partner only. In the responding pairs, there were highly significant increases in mounting attempts and ejaculations, an effect that could be attributed only to treatment. We conclude, therefore, that these aliphatic acids (copulins), which act via olfactory pathways, have communicatory significance in rhesus monkeys.     

Behaviorial effects of a synthetic mixture of aliphatic acids in rhesus monkeys (Macaca mulatta).

The stimulation of sexual behavior by a synthetic mixture of volatile aliphatic acids (acetic, propanoic, methylpropanoic, butanoic, methylbutanoic, methylpentanoic) was studied in male rhesus monkeys. Twelve intact adult males and 12 long-ovariectomized adult females were used in 24 paired combinations (541 tests each of 1 hr). A mixture of authentic acids similar to that found in the vaginal secretions of estrogenized females was applied to the sexual skin area of ovariectomized females immediately before tests with males. There was marked between-pair variability during the application of both control and test substances. However, using rigorous behavioral criteria, there was a well-marked stimulation either of male mounting attempts or of ejaculations in 12 of 24 pairs involving 9 of 12 males. Three males responded with both female partners, three responded with neither female partner, and six responded with one partner only. In the responding pairs, there were highly significant increases in mounting attempts and ejaculations, an effect that could be attributed only to treatment. We conclude, therefore, that these aliphatic acids (copulins), which act via olfactory pathways, have communicatory significance in rhesus monkeys.     

A review of sexual initiating behavior by male and female cynomolgus monkeys and some species comparisons

The sexual initiating behavior of male and female cynomolgus monkeys (Macaca fascicularis) observed during standard laboratory tests is reviewed and compared with that of rhesus monkeys (M. mulatta) observed under identical conditions. Species differences in sexual behavior are related here to differences in habitat, sexual dimorphism, and the dominance gradient between the sexes. Compared with rhesus monkeys, cynomolgus monkeys appear to be more arboreal, less sexually dimorphic, and have a smaller dominance gradient between the sexes. They exhibit a facultative single-mount copulatory pattern rather than the serial mount pattern of the rhesus monkey. Female cynomolgus monkeys are less dominated than rhesus females by their male partners. Direct aggression between mates is more frequent and redirected aggression occurs less often than in rhesus monkeys. These behavioral differences affect the interpretation of changes in initiation rates that occur (1) during the menstrual cycle, (2) when females are ovariectomized and given hormone replacement treatments, and (3) when males are castrated and treated with androgens. We conclude that estradiol in the female and testosterone in the male increase the sexual motivation of both the treated and the untreated partner. Valid interpretations of changes in initiation rates depend on accurate and exclusive definitions of behavior and on a consideration of the behavioral context in which they are made.     

Female dominance rank and behavior during artificial menstrual cycles in social groups of rhesus monkeys (Macaca mulatta)

We recently reported (Michael & Zumpe: American Journal of Primatology 15:157–170, 1988) evidence for the hypothesis that copulations by pairs of rhesus monkeys are linked more closely to the ovulatory phase of the female's menstrual cycle when a male has access to several females in different cycle phases rather than access to females in the same cycle phase. We have now used data from this earlier study to assess the role of female dominance. Artificial menstrual cycles were induced with hormone injections in two of four ovariectomized females in eight social groups (each consisting of one male and four females). The artificial cycles were either synchronized or offset by 7-day increments. There were differences in the behavioral interactions of the higher-ranking (dominant) and lower-ranking (subordinate) hormone-treated females in each group. The amount of male sexual activity received by dominant and subordinate females differed little, but dominant females initiated significantly more sexual activity (P<.0001) than did subordinate females. Dominant females were significantly more aggressive (P<.0001) toward other females and received more direct and redirected aggression from males. Males generally initiated and maintained more social contacts with dominant than with subordinate females, and dominant females were less active than subordinates in initiating and maintaining social interactions with males. The findings provide new evidence for both female mate competition and mate choice by males for higher-ranking females.     

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.     

Sex-specific interactions between aggressive and sexual behavior in the rat: Effects of testosterone and progesterone

The influence of progesterone on sexual and aggressive behaviors during aggressive encounters was investigated in pairs of TP-treated male and female rats. Gonadectomized females, chronically injected with testosterone propionate (TP), showed low but consistent levels of feminine sexual behavior which alternated with aggression. Progesterone when given in addition to TP facilitated receptive and proceptive behaviors, but reduced levels of aggression. In TP-treated males, levels of aggression were the same as observed in TP-treated females. However, TP-treated males seldomly showed sexual behavior during aggressive encounters and additional treatment with progesterone did not affect their behavior. After the aggression tests, animals were tested in a social preference test in which an ovariectomized female cage mate and the opponent from the aggressive encounter served as incentives. Positive correlations between levels of aggression and social preference for an opponent were found in both sexes, although correlations only reached statistical significance when progesterone was given in addition to TP. These correlations were found in both sexes, despite the fact that group analysis revealed pronounced sex differences in social preference: males preferred to spend their time near ovariectomized female cage mates, whereas females divided their time equally among female cage mates and opponents.     

Does Female Personality Determine Mate Choice Through Sexual Cannibalism?

Animal personalities (e.g. consistent across‐context behavioural differences between individuals) can lead to differences in mate choice. However, evidence for this link remains limited. Pre‐mating sexual cannibalism can be a behavioural syndrome (i.e. a suboptimal personality) in which adaptive female aggression towards heterospecific prey spills over on non‐adaptive aggression towards courting males, independently of the female mating or feeding status (i.e. the ‘aggressive spillover hypothesis’, ASH). On the other hand, sexual cannibalism can also be a form of mate choice by which females selectively kill or mate with males depending on the male phenotype. We introduce the hypothesis that the most aggressive females in the population will not only attack males more frequently, but will be less likely to impose sexual selection on males through sexual cannibalism. Assuming that in a field common garden experiment in which females were fed ad libitum the rate of weight gain by a female may reflect her voracity or aggressiveness, we show that in the cannibalistic burrowing wolf spider Lycosa hispanica (formerly L. tarantula), voracity towards heterospecific prey predicts a female's tendency towards sexual cannibalism. Unmated females with higher weight gains were more cannibalistic and attacked males regardless of the male phenotype. On the other hand, females that were less voracious tended to be less cannibalistic, and when they did kill a male, they were selective, killing males in poorer condition and mating with those in better condition. Our results demonstrate that females with different phenotypes (growth rates) differently imposed selection on male condition, tentatively supporting the hypothesis that female aggression levels can spill over on sexual selection through sexual cannibalism.     

Sociality in Callithrix penicillata: II. Individual Strategies During Intergroup Encounters

In social animals, intergroup interactions, whether through agonistic and competitive behaviors or affiliative ones, can influence important parameters such as home range, territory sizes, and access to resources, which may directly affect both female and male fitness. We studied the intergroup interaction patterns of a wild group of black-tufted-ear marmosets (Callithrix penicillata) in central Brazil. Agonistic interactions occurred at low frequencies during intergroup encounters. The marmosets directed agonistic interactions without physical aggression primarily against same-sex individuals, suggesting that male and female aggression patterns are shaped by their sexual interests. However, females of the focal group also directed agonistic behavior toward extragroup males that attempted copulation. The marmosets appeared to use intergroup encounters to gather information about possible partners and extragroup reproductive opportunities. Intergroup sexual interactions occurred mainly in the form of copulations or attempted copulations by all adults, with the exception of the dominant female. Our results suggest that a possible reproductive strategy used by males is to attempt fertilization of extragroup females. Adult males copulated with the same extragroup female during several opportunities, which suggests sperm competition or the establishment of social bonds with neighboring females.     

Mating Tactics in Response to Costs Incurred by Mating with Multiple Males in Wild Female Japanese Macaques

We investigated the costs of mating with multiple males in terms of feeding time, traveling distances, sexual proceptivity, and male aggression, for wild female (Macaca fuscata yakui) on Yakushima Island, Japan. We analyzed all-day focal sampling data from 7 females during the mating season (Sept.-Nov. 1996). On days when estrous females copulated with multiple males, they decreased their feeding time to half that of anestrous days, traveled longer distances, showed more proceptive sexual behaviors and received more aggression from subordinate males than on days when they copulated with only the 1st-ranking male. On days when females copulated with only the 1st-ranking male, they showed no difference in feeding time with that of anestrous days, and expended less effort than the above mating pattern because of short traveling distances, diminished sexual proceptivity and a lower frequency of aggression received. The results suggest that the costs of estrous vary according to female sexual proceptivity and the number and social status of mating partners. Female Japanese macaques exhibit a mixed mating strategy over prolonged estrous periods, which may provide females with opportunities to maximize the benefits of copulating with multiple males and to minimize the costs of estrus by mating with only the 1st-ranking male. During an estrous cycle, females may be adjusting efforts for reproduction and survival; i.e., mating vs. feeding.     

Temperament in a Fish: A Longitudinal Study of the Development of Individual Differences in Aggression and Social Rank in the Midas Cichlid

A longitudinal design was applied in a study of the development of individual differences in aggression, and their relationship to growth and sexual differentiation in the Midas cichlid, Cichlasoma citrinellum. Weight and standard length measurements, as well as several measures of aggression, were obtained at regular intervals. Size ranks within groups were stable over a period spanning an early juvenile phase through sexual maturity. Two of three aggression rank scores were also stable over this interval, but they did not relate in any direct way to social rank. As juveniles, males showed higher levels of aggression than females, but the reverse was true in adults. This sex-specific developmental change resulted both from an increase in female aggression at sexual maturity and a decline in male aggression at sexual maturity. Individuals showed distinct and stable behavioral profiles with respect to the combined aggression tests.     

Influence of the estrous cycle on heterosexual aggression in two strains of rats (S3 and WEzob)

Two strains of rats (S₃ and WEzob), which show different levels of aggression in the laboratory, were tested in repeated heterosexual confrontations. Daily 15-min observations were made of the interactions between a female throughout a complete estrous cycle and the same male partner. In both strains the topography of aggression was similar in males and females, but the frequency of specific parameters varied. Males showed more offensive and females more defensive patterns. The overall level of aggression was very low on the day of estrous, when the female was sexually receptive. There were no differences in any elements of female or male behavior between the other 3 days of the cycle. The results support previous conclusions from single-sex encounters that in rats there is no sexual dimorphism in the ability to show aggression.     

Ontogenetic changes and the stability of rhesus monkey dominance relationships

Dominance relationships were studied in a rhesus monkey group during five consecutive years. The group consisted of eight stable matriarchies and an adult male class which was replaced at the start, and again at the midpoint, of the study. Immature males were selectively harvested to maintain a sex ratio typical of natural troops. Maximum group size during the study was 77 animals.Dominance relationships were remarkably stable, with only 4.4% of dyads failing to show unidirectional relationships. Despite this stability, a linear ranking of all group members was not possible. Male dominance relationships with other males were among the most stable, following the fighting which ensued on male introductions. Male introductions did not disrupt female dominance relationships.Adult female dominance relationships were also quite stable, but immature females slowly achieved dominance over older sisters and females subordinate to their mothers. Such reversals were the result of processes lasting over many months. Many dominance assertions occurred prior to puberty but a significant number occurred following sexual maturity. Maturing females did not reverse dominance relationships according to any particular hierarchial order and, as a consequence, many were subordinate to animals that were dominated by others that they dominated.Although there was an alpha male that was dominant to all animals in the group, adult females dominated most adult males. Adult males, however, often reciprocated aggression directed at them. They almost invariably threatened or countercharged aggressive immature animals regardless of matriarchial membership. Adult males dominated some adult and most young females, even in families containing matriarchs and adult females to which the adult males always submitted.The dominance relationships of young males were similar to those of their sisters, until puberty. Young males did not necessarily bypass adult males that their mothers outranked, and often failed to win against adult females that their mothers dominated. Adolescent female aggression against females is seldom interfered with by adult males, and females may actively aid one another against males. In contrast, the aggression of young males often elicits interference by adult males, and young males often become the targets of redirected aggression in the group. As a consequence, whereas young females rise in rank to positions adjacent to their mothers, adolescent males often suffer losses to animals that they had dominated as juveniles.     

Parity of female goats does not influence their estrous and ovulatory responses to the male effect

The objective of the present study was to determine whether parity is a factor that influences the estrous and ovulatory responses of female goats when they are stimulated by males that show increased sexual activity. To stimulate sexual activity, four adult male goats were subjected to photoperiodic treatment for 2.5 months comprising long days, with the treatment commencing on 1 November. On 14 April at 1900 h, a group of multiparous females (n = 21) and a group of 16 months-old nulliparous females (n = 19) were exposed to four bucks (two per group) for 15 days. Throughout the study period, the estrous behavior of these female goats was detected twice on a daily basis. Ovulations of the female goats were determined by ecography on days 7 and 18 after exposure to males. The sexual behavior of males was recorded twice every day from 0800 to 0900 h and from 1730 to 1830 h during the first 4 days after introduction in the pen of females. The total cumulative proportion of multiparous females that had ovulations (100%) and displayed estrous behavior (100%) during the 15 days of exposure to males did not differ (P > 0.05) from that of nulliparous females (100% and 95%, respectively). The interval between introduction of males and onset of estrous behavior did not differ (P > 0.05) between multiparous (1.9 ± 0.1 days) and nulliparous (1.7 ± 0.2 days) females. The proportion of females displaying a short estrous cycle was greater (P < 0.05) in multiparous (13/21, 62%) than in nulliparous (5/19, 26%) females. Duration of these shorter than typical estrous cycles did not differ (P > 0.05) between groups (multiparous: 5.2 ± 0.3 days, nulliparous: 4.5 ± 0.1 days). The number of anogenital sniffings was greater (P < 0.001) in males exposed to nulliparous than in those exposed to multiparous females. In contrast, the number of mounting attempts was greater (P < 0.01) in males that were introduced to multiparous than in those that were introduced to nulliparous does. The number of flehmen, nudging, self-marking with urine, and mounts was not different (P > 0.05) between males that were in contact with multiparous and nulliparous females. These results indicate that regardless of parity, female goats respond to male introduction if they are stimulated by males that were previously exposed to artificial long days to increase their sexual behavior.     

Female genital swellings have no effect on chimpanzee agonism in well-established groups

Reducing the incidence of aggression and wounding is a major concern for those managing socially housed chimpanzees (Pan troglodytes). Competition among adult males for access to sexually receptive females is believed to be one factor contributing to undesirably high levels of agonism. An observational analysis of two chimpanzee groups (n = 9, n = 8) was initiated to study this phenomenon. All-events sampling was used to record agonistic and sexual behavior on days when one female in the group showed a maximal genital swelling and on other days when no female in the group had a tumescent swelling. The result of the multivariate analysis of variance indicated that the presence of a female with a tumescent swelling had no effect on bluff displays or aggressive, submissive, or reconciliatory behavior but that sexual behavior was significantly increased. These results may be explained by the fact that long-term stability in group membership may reduce male–male competition for sexual access to females and the sometimes associated aggression and wounding. Colony managers dealing with well-established groups of chimpanzees should invest more of their effort in controlling aggression elicited by factors other than the cycling of a single female group member.     

Breeding Season Aggression of Female and Male Eastern Bluebirds (Sialia sialis) to Models of Potential Conspecific and Interspecific Egg Dumpers

Naturally occurring aggression between female eastern bluebirds (Sialia sialis) is dramatic, resulting in severe injuries and even death. Furthermore, aggression among bluebirds is usually sex specific: males attack males, females attack females. We hypothesized that the primary function of female-female aggression is to guard against the threat of intraspecific egg dumping and that, in this context, same sex aggression is related to the possibility of advantages for males of parasitism (egg dumping) of their nests. Our hypotheses to explain variation in naturally occurring aggression predict temporal variation in aggressive tendency within nest cycles and between the sexes depending on asymmetries in threats to the residents. We report the results of experimental trials in the field designed to determine temporal variation in the aggressive tendencies of resident females to models of intruder females of two species, eastern bluebirds and brown-headed cowbirds (Molothrus ater). Both species dump eggs in the nests of bluebirds. Female aggression to eastern bluebird models is greatest during early stages of nest cycles; the patterns are most consistent with protection against egg dumping and protection of nest sites from usurpation. Male residents seldom attack female eastern bluebird models, but often attack models of female brown-headed cowbirds, a result inconsistent with the hypothesis that patterns of differential parental care control aggression of female and male residents.     

Interactions of social,spatial and hormonal factors on the behavior of rhesus monkeys (Macaca mulatta)

In a three year study, the interactions of social, spatial and hormonal variables on the behavior of rhesus monkeys were investigated. Observations were made on (1) an oppositelysexed pair alone in a cage; (2) the pair alone in a large room; and (3) the pair together with three ovariectomized females in the same large room. Each of four males was tested with the same group of four ovariectomized females (16 pairs, 480 1-hr tests). Each female served in turn as the focal female, namely, the hormone-treated one, and received first estradiol benzoate (EB) alone and then in combination with progesterone (P). The presence of three other females (the social variable) had a marked effect on all aspects of the behavior of the focal pair, and reduced the male's ejaculations as well as other sexual and affiliative interactions with the focal female. This was due to an inhibitory influence exerted by the dominant female on the interactions of subordinate females with the male. Comparing data from two groups of females showed that female dominance rank, and not male sexual preference, was the critical factor, and that mate competition between female rhesus monkeys was operating. Decreasing the space available for the interactions of the pair appeared to imitate a bonding effect in the male, increasing male grooming and tending to increase male sexual activity. In the group setting, treating the female with EB increased sexual activity, and additional progesterone decreased it in all three test settings. The effects of progesterone were most clearly apparent when the pair was tested alone in a large area.     

Gender differences in the risk of HIV infection among persons reporting abstinence, monogamy, and multiple sexual partners in northern Tanzania

Background

Monogamy, together with abstinence, partner reduction, and condom use, is widely advocated as a key behavioral strategy to prevent HIV infection in sub-Saharan Africa. We examined the association between the number of sexual partners and the risk of HIV seropositivity among men and women presenting for HIV voluntary counseling and testing (VCT) in northern Tanzania.

Methodology/ Principal Findings

Clients presenting for HIV VCT at a community-based AIDS service organization in Moshi, Tanzania were surveyed between November 2003 and December 2007. Data on sociodemographic characteristics, reasons for testing, sexual behaviors, and symptoms were collected. Men and women were categorized by number of lifetime sexual partners, and rates of seropositivity were reported by category. Factors associated with HIV seropositivity among monogamous males and females were identified by a multivariate logistic regression model. Of 6,549 clients, 3,607 (55%) were female, and the median age was 30 years (IQR 24–40). 939 (25%) females and 293 (10%) males (p<0.0001) were HIV seropositive. Among 1,244 (34%) monogamous females and 423 (14%) monogamous males, the risk of HIV infection was 19% and 4%, respectively (p<0.0001). The risk increased monotonically with additional partners up to 45% (p<0.001) and 15% (p<0.001) for women and men, respectively with 5 or more partners. In multivariate analysis, HIV seropositivity among monogamous women was most strongly associated with age (p<0.0001), lower education (p<0.004), and reporting a partner with other partners (p = 0.015). Only age was a significant risk factor for monogamous men (p = 0.0004).

Interpretation

Among women presenting for VCT, the number of partners is strongly associated with rates of seropositivity; however, even women reporting lifetime monogamy have a high risk for HIV infection. Partner reduction should be coupled with efforts to place tools in the hands of sexually active women to reduce their risk of contracting HIV.     

Female-Directed Aggression in Free-Ranging Ateles geoffroyi

In a study of the reproductive biology and behavior of black-handed spider monkeys (Ateles geoffroyi) on Barro Colorado Island, Panama, 107 instances of female directed male aggression (male attacks) occurred during >1,000 h of observation. Two hypotheses might explain this behavior. First, males may attack to induce defecation/urination by target females, from which they can gain reproductive information. Rates of defecation and urination did not increase subsequent to attacks; however, there was an association between attacks and investigative behaviors such as place sniffing. Alternatively, the attacks might be a form of sexual coercion. My results indicate that female reproductive state, as determined by fecal steroid metabolites (E1C and PdG), had an inconsistent effect on the frequency of attacks received. In addition, when females were in the peri-ovulatory period of the ovarian cycle they did not receive more attacks than at other times. Nor were attacks associated with observed copulations, suggesting that sexual coercion does not sufficiently explain the behavior. Some support is provided for the notion that the attacks allow males to dominate equally-sized females.