The behavior of peripheral males during the mating season inMacaca fuscata

The mating season behavior of peripheral male Japanese macaques (Macaca fuscata) at Arashiyama West were studied during the 1994/1995 mating season. Although all peripheral males increased their proximity to the main troop, there was great variation in this behavior, from those who became virtually indistinguishable from the main troop males to those who moved in and out of the main troop in a clandestine fashion, to those who had only visual contact, from a distance of 25 m, with the main troop. The subsequent behavior of males displaying these patterns was compared, as was the behavior of peripheral to main troop males. Specifically, they were investigated for variation in three behavioral strategies which may function to increase access to mates: aggressive intimidation; affiliation with unrelated main troop females; and courtship display. The results show that peripheral males are a more diverse group than previously described, and that one group of peripheral males was able to successfully join the main troop, becoming indistinguishable from main troop males with regard to these behaviors which enhance access to mates.     

Socio-sexual factors of troop fission in wild Japanese monkeys (Macaca fuscata yakui) on Yakushima Island,Japan

The troop fissions which occurred in a wild population of Japanese monkeys (Macaca fuscata yakui) were observed from 1977 to 1979 on Yakushima Island. The fissions were initiated in the breeding season by non-troop males who established a consort relation with estrous females. In order to analyze the socio-sexual factors which accelerated the fissions, the male emigrations and immigrations before and after two successive fissions, and the copulation frequencies, competition among males and preferences of mating partners in both sexes in the 1977–78 breeding season after the first fission were examined. The results indicated that three factors (a large number of non-troop males, a shortage of troop males and the females' choice of mating partners) effectively influenced on the establishment of consort relationships between non-troop males and estrous females. It is suggested that these factors may exert different effects on the troop disorganization in relation to troop size. In small-sized troops, a large number of non-troop males and a shortage of troop males may lead to stronger competition between them, and the females' choice affected by prolonged intimate relations with the dominant TMs may reduce their priority of access to estrous females. This situation possibly stimulates fission or male emigration in small-sized troops under the natural conditions on Yakushima Island. In contrast, in large-sized troops under isolated conditions, a surplus rather than a shortage of troop males may contribute to troop disorganization, as most former studies have suggested. A higher socionomic sex ratio may decrease the mating activities of subordinate troop males and increase the competition among them. This situation possibly accelerates the fission of large-sized troops through prolonged interactions between females and subordinate or peripheral troop males. A lower ratio and the females' choice, however, raise the mating chances of subordinate troop males and may not promote the fission of large-sized troops under isolated conditions. This study was financed in part by a Grant-in-Aid for Special Project Research on Biological Aspects of Optimal Strategy and Social Structure from the Japan Ministry of Education, Science and Culture, and by the Cooperative Research Fund of the Primate Research Institute, Kyoto University.     

Affiliative relations among male Japanese macaques (Macaca fuscata yakui) within and outside a troop on Yakushima Island

Male Japanese macaques (Macaca fuscata yakui) in a troop on Yakushima Island frequently groom other males. However, previous studies have not compared the social relations of troop males to those of non-troop males. I followed all troop males and non-troop males in and near a troop during a mating season and during the following non-mating season and recorded their neighbors, grooming, and agonistic interactions. Comparisons of the social relations of troop males and non-troop males with other troop members revealed that grooming and agonistic interactions with females during the mating season were similar between troop and non-troop males. However, troop males groomed each other more often and had fewer agonistic interactions among themselves than did non-troop males. Compared to what occurred in the mating season, troop males groomed females less often and exchanged grooming bouts more often with other troop males during the non-mating season. One non-troop male groomed females more frequently than did any troop male in both seasons, and this male groomed troop males more frequently than did any troop male in the non-mating season. This male immigrated into the troop during the following mating season. Regardless of their competition with respect to reproduction, male Japanese macaques on Yakushima Island maintain affiliative relations, probably to cooperatively defend fertile females from non-troop males.     

Hormonal Correlates of Social Rank in an Asocial Species, the Common Brushtail Possum (Trichosurus vulpecula)

Dominance hierarchies play an important role in access to mates or resources in many species. Rank is sometimes correlated with circulating testosterone levels or morphological traits such as body weight. The relationship of glucocorticoid secretion and rank, however, is less clear. In this study, we investigated the relationship of male rank to body weight, circulating testosterone and cortisol concentrations in captive possum triads (Trichosurus vulpecula). We carried out two experiments to examine hierarchy formation and the effects of castration of the dominant male during the non‐breeding season. A third experiment measured the effects of removal of the dominant male from a stable hierarchy during the breeding season. We found that dominant male rank was significantly correlated with higher circulating testosterone levels during periods of hierarchy formation and during the breeding season but not during periods of hierarchy stability in the non‐breeding season. Lack of correlation between plasma testosterone concentration and rank after male castration suggests that stable social rank is not dependent on hormone level and may be more dependent on behavioural traits. Any biocontrol measures that rely on manipulation of hormone levels may be unreliable when applied to unstable hierarchical situations, including the establishment of territories by subadult males, and the pre‐breeding season when circulating testosterone concentrations peak in males and a period of hierarchy establishment may occur.     

Effects of social status, age, and season on androgen and cortisol levels in wild male golden lion tamarins (Leontopithecus rosalia)

This study examines factors affecting androgen and cortisol levels in wild, male golden lion tamarins (Leontopithecus rosalia). Golden lion tamarins are a cooperatively breeding species in which groups often contain two potentially breeding males. Brothers frequently emigrate together and develop a clear dominance hierarchy, but interactions between them are primarily affiliative. Duos in which the males are not related are less stable. In addition, reproductive skew theory predicts that dominant males will be less likely to share reproduction with related subordinates. As such, we predicted that both androgens and cortisol would be higher in subordinate males unrelated to the dominant male. We also predicted that androgens in breeding males would be higher during the mating season than the birth/infant care season, as per Wingfield's "challenge hypothesis" (1990). Fecal samples were collected from 24 males in 14 social groups and assayed by enzyme immunoassay. Androgen levels were higher in breeding males during the mating season, thus supporting the challenge hypothesis. However, while subordinate males unrelated to the dominant male had significantly lower androgens than any other group, cortisol levels were not correspondingly higher. These results suggest that unrelated subordinate males show measurable reproductive suppression and may use strategies such as infantilization to avert aggression from dominant males.     

Seasonal variation in sexual behavior of all-male rhesus troops

Free roaming groups of Rhesus monkeys show a clear annual cycle in the incidence of sexual behavior. One hypothesis explaining the control of this cycle is that seasonal variables act directly on females, altering their endocrine status. Aroused females then communicate their sexual receptivity to male group members. The sexual behavior of 35 male Rhesus was observed during the non-mating and the mating season. All animals were maintained in large outdoor enclosures. One group of males (N = 7) were housed with females and juveniles as part of a breeding group. A second group of males (N = 8) had visual contact with the members of the breeding group, and a third group of males (N = 20) had no contact with either of these groups. During the non-mating season no sexual behavior was observed. At the onset of the mating season, the males with females displayed evidence of hormonal arousal and became sexually active. The males with visual contact with the breeding group also evidenced hormonal arousal and began engaging in male-to-male sexual activity. The males in the third group showed no sign of sexual arousal. These findings support the hypothesis that seasonal influences act only on females; and demonstrate that females can communicate their endocrine status to males, even without direct physical contact.     

Behavioural and hormonal responses of male rhesus monkeys introduced to females in the breeding and non-breeding seasons.

Six adult male rhesus monkeys were introduced individually to an all-female group for 10 days during the mating season. The initial aggressive responses of the females were rapidly replaced by positive social behaviour, and each male achieved alpha status and had access to social and sexual partners. A repetition of this paradigm in the non-breeding season produced significantly more female aggression, and no male attained high rank or engaged in sexual or other social behaviour. Male testosterone levels rose following introduction to the females in both seasons, but were significantly higher during the breeding season. Hormonal levels following removal from the females suggest a complex interplay between social, sexual and seasonal variables and recent social experiences. The differences in female social behaviour with newly introduced males, as a function of season, suggest an explanation for the seasonal limitation of male troop transfers.     

Display behavior of three troops of Japanese monkeys (Macaca fuscata)

In this paper the display (i.e., “branch-shaking”) behavior of three troops of Japanese monkeys—the confined Oregon troop as reported byModahl andEaton (1977), the semi-free-ranging Arashiyama West troop and the free-ranging Arashiyama B troop—are compared. The comparison reveals several similarities and dissimilarities between the Oregon troop and the two genetically related Arashiyama troops. The similarities include three display postures (shaking, kicking and leaping) and an increased frequency of male but not female displaying during the breeding season. The dissimilarities include the absence of two Oregon display postures (tossing and swinging) and collective displaying among Arashiyama monkeys. The hypothesis ofModahl andEaton (1977) that male displaying influences the females' choice of male mating partners is discussed. The evidence suggests the Oregon monkeys have incorporated male displaying into their socio-sexual behavior to a greater extent than Arashiyama monkeys. However, field observations indicate that males who migrate to the Arashiyama B site during the breeding season can influence female choice by “advertizing” their location with displays.     

Why should an aged male baboon ever transfer troops?

Dispersals by subadult or adult male baboons are common. In contrast, the occasional transfer of aged baboons is puzzling, given the physical dangers of transfer and the cognitive demands of mastering the ecological and social rules of a new troop. The present data suggest a possible explanation for such transfers in a study of two troops of olive baboons (Papio anubis) in Kenya. Aged males who remained in the troop in which they had been dominant were subjected to significantly higher rates of approach-avoid interactions by the current high-ranking cohort (i.e., the individuals they had dominated years past) when compared to males who had transferred into the troop in their old age. Thus, transfer in old age offers the advantage of relative anonymity. Of the 14 males who progressed into older adulthood in the same troop in which they were in their prime, seven ultimately transferred to a different troop. The seven who remained and the seven who transferred did not differ in the rate at which they were subjected to approach-avoid interactions. However, those who remained had significantly higher rates of various affiliative behaviors (copulations, consortships, grooming and contact with females, and positive interactions with infants). Thus, amid the disadvantages of an old age spent in the troop in which a male baboon was in his prime, a high degree of social affiliation might constitute a sufficient disincentive against transferring. © 1996 Wiley-Liss, Inc.     

Social organization of wild talapoin monkeys

A troop of wild talapoins in Cameroon had 70 members including all age-sex classes. Observed between the birth season and the next mating season, it was habitually divided into subgroups. Subgroups most commonly observed were (1) adult and large juvenile males: (2) adult females with infants, small juveniles, and large juvenile females; (3) medium sized juveniles with a single adult male. Adult females and adult males were rarely sighted together and never seen to interact. From observations of caged groups, predictions are made about the changes in this organization which might be expected during the breeding season. A possible parallel with the ecologically similar squirrel monkey is suggested.     

Factors Influencing the Breeding Activity in Male Masu Salmon, Oncorhynchus masou: The Relationship with Body Size

Synopsis Male body size has been emphasized as an important factor contributing to the breeding success of individuals. However, the operational sex ratio (OSR: ratio of mature females to males) during the breeding season significantly change due to differences in the breeding timing and period within and between males and females and may influence the selective advantage of the male large body size for mates. We examined the reproductive ecology of masu salmon Oncorhynchus masou, inhabiting Lake Toya, Hokkaido, northern Japan. Precipitation triggered the upstream migration, although males migrated into the river earlier than females. As a result, the OSR in the river changed markedly during the breeding season, as did the size structure of males in the river. Large males migrated into the river earlier than the smaller males. Differential male survival resulted in the different population size structure between the early and late periods. Under these conditions, we analyzed which males were more successful in releasing the more sperm throughout the breeding season by estimating the decrease in the sperm content in male carcasses. The body size and sperm-releasing success of the males were not related. With a weak male-biased OSR and synchronous timing of reproduction in females, males that had entered the river succeeded in releasing the sperm regardless of body size. Such a fluctuating advantage for different body sizes likely contributes to the maintenance of the size variation in the male salmon.     

Age,condition and dominance‐related sexual ornament size before and during the breeding season in the black grouse Lyrurus tetrix

Male ornaments function as honest cues of male quality in many species and are subject to intra‐ and intersexual selection. These ornaments are generally studied during peak expression, however their size outside the breeding season may determine ultimate ornament size and costliness, and as such reproductive success. We investigated whether male black grouse Lyrurus tetrix eye comb size was related to age, condition and measures of male dominance before and during the breeding season. Total combined eye comb size began to increase ~70 d before the start of the breeding season. Adult males (aged ≥ 2 yr old) had consistently larger eye combs than younger males (1 yr old) both before and during the breeding season. Heavier and more dominant adult males (attending the lek more frequently and successfully reproducing) had larger eye combs. For younger males, those that were heavier had larger eye combs. Additionally, males that spent more time on the lek showed increased eye comb size as the breeding season approached. Overall we find that ornament size is positively related to dominance and condition before and during the breeding season. Since dominance is accrued through year‐round interactions in many species, the ability to maintain larger signals over prolonged periods, including outside of the breeding season, is likely to be beneficial for adults. For younger males, it is likely that they cannot sustain or are constrained from producing larger eye combs over long periods of time. They therefore prioritise growth of their ornaments later, and according to the amount of time they spend on the lek.     

THE BREEDING CYCLE OF THE SOUTHERN SEA LION, OTARIA BYRONIA

Daily counts of Southern sea lions ( Otaria byronia ) made at Punta Norte, Argentina, during four consecutive breeding seasons (1980–1984) yielded similar results in the time of the reproductive events. Males and females began to arrive during the second week of December. Males reached peak numbers (100–110) between 15 and 21 January. Females reached maximum numbers (300–350) at the end of January. About 380–430 pups were born between December 20 and February 2, and 70–80 percent of births occurred between 10 and 25 January. Copulations peaked during the third week of January. The sex ratio of adult males to adult females at peak season fluctuated from 1:3.7 to 1:4.2. By the first week of February, coinciding with the maximum number of young males (25–50), 90 percent of the adult males had abandoned the area and the number of females fluctuated greatly. Since 1980 the number of females and pups has shown a slight increase, particularly during the 1983–1984 breeding season.     

Resting cortisol levels and the emergence of dominant status among male vervet monkeys

Resting serum cortisol was measured in adult male vervet monkeys (Cercopithicus aethiops sabaeus) in four different conditions: (1) among groups with unaltered group membership and established dominance hierarchies; (2) among groups from which the original dominant male had been removed and in which the remaining males competed for dominant status; (3) among newly formed groups of three unfamiliar males each of which had been the dominant male in his previous group; and (4) among groups from which a dominant male was temporarily separated and returned. In Condition 1, cortisol concentrations did not differ between dominant and subordinate males. The second condition showed that cortisol levels were highest among males who eventually emerged as the dominant male. In the third condition, however, cortisol levels did not differentiate eventually dominant from eventually subordinate males. In the last condition, cortisol levels were highest in the animals that became or remained dominant following reintroduction. These data indicate that cortisol concentration does not differ between dominant and subordinate males in stable groups and that cortisol rises during competition for dominance among familiar males.     

A longitudinal analysis of reproductive skew in male rhesus macaques

One of the basic tenets of sexual selection is that male reproductive success should be large in polygynous species. Here, we analysed 6 years of molecular genetic data from a semi-free-ranging population of rhesus macaques (Macaca mulatta), using Nonac's B index, to assess the level of male reproductive skew in the study troop. On average, the top sire in each year produced 24% of the infants, while 71% of troop males sired no offspring at all. Consequently, 74% of infants had at least one paternal half-sibling in their own birth cohort. Reproductive success was greatest for high-ranking males, males who spent the whole mating season in the troop and males of 9-11 years of age. Heterozygosity for major histocompatibility complex (MHC) class II gene DQB1 was the strongest single predictor of male reproductive success. A negative relationship suggestive of female mate choice was noted between the B index and the proportion of extragroup paternities. Reproductive skew was not associated with relatedness among potential sires or with female cycle synchrony. We conclude that reproductive skew in male rhesus macaques is best accounted for by the 'limited-control' model, with multiple factors interacting to regulate individual reproductive output.     

Seasonal changes of serum sex steroids concentration and aromatase activity of gonad and brain in red-spotted grouper (Epinephelus akaara)

Levels of serum sex steroids (estradiol-17beta, E2; testosterone, T; 11-ketotestosterone, 11-KT) in male, female and natural sex-reversing red-spotted grouper (Epinephelus akaara), and aromatase activity of gonad and brain in both male and female were investigated throughout an annually reproductive cycle. In females, serum E2 and T peaked during vitellogenesis, but in males and natural sex-reversing fish, 11-KT, T and E2 reached peak during spermatogenesis. In addition, in females, serum 11-KT levels (monthly means: 0.32 +/- 0.03 ng/ml) which were very low did not significantly fluctuate during the annual reproductive cycle. In breeding season, females displayed higher E2 levels than males and sex-reversing fish, while males and sex-reversing fish showed higher 11-KT levels and, to a lesser extent, higher T levels than females. Furthermore, the changing pattern of sex steroids in males was similar to that in natural sex-reversing fish, and a second peak of serum androgens 11-KT and T appeared in December both in male and natural sex-reversing fish; significantly higher serum 11-KT levels were observed in natural sex-reversing fish than that in females from December to April. In females, but not in males, aromatase activity of brain and gonad demonstrated significantly seasonal changes (exhibiting a peak in breeding season); moreover, aromatase activity in females was higher than that in males. Furthermore, significantly lower aromatase activity in testis was observed in breeding season, in contrast to that in ovary. Taken together, the present findings indicated that changes of serum sex steroids levels and aromatase activity in red-spotted grouper were closely associated with sex inversion. In addition, the present results also suggested that sex inversion in red-spotted grouper peaked mainly from December to March.     

Male competition and coalitions in langurs (Presbytis entellus) at Jodhpur,Rajasthan, India

During a 15 month study on free ranging langurs (Presbytis entellus) at Jodhpur, Rajasthan, India, 5 adult male replacements were observed as a result of nontroop male invasions into the home ranges of 3 neighbouring one-male troops comprising 16–28 members each. Jodhpur langurs have no breeding season. Periods of instability during resident male changes lasted 11–119 days. Linear dominance hierarchies could be detected within the 3 main rival male bands of 2, 5, and 28–35 members. The respective alphas drove their allies away after their bands succeeded cooperatively at occupying a troop. During gradual replacements interim residencies alternated with multi-male stages. A large band's alpha may have had better chances to win the competition, since adult and nonadult allies functioned as “buffers” in agonistic encounters. The role of kin selection in structuring the composition of male bands and male coalitional behaviour cannot yet be quantified. Tactical “deceit” of powerful males to cause unrealistic expectations and in this way agonistic engagement of less strong males can be ruled out. “Sneaking copulations” is a proximate advantage for subordinate supporters, since they participated in 61.9% of all sexual interactions. Female promiscuity might reflect a strategy to induce male-male competition and thus select for a strong resident.     

The social organization of forest hanuman langurs(Presbytis entellus)

The social organization of hanuman langurs (Presbytis entellus;Colobinae) was studied in Kanha Tiger Reserve, Central Indian Highlands, for 2300 hr (1980–1985), in a mosaic of moist deciduous forest and anthropogenic meadow. The langur population density was 46.15/km ² and the mean troop and band sizes were 21.7 and 14.0, respectively. Of 14 troops, 13 were one-male and 1 was trimale. The population adult sex ratio was 1:2.5. The majority of female sexual solicitations was directed toward the harem male. The birth season was December to May, with an estimated gestation of 171–224 days. A review of langur reproductive seasonality suggests that breeding throughout the year is confined to those populations able to exploit human food sources. Mortality during the first year of life was 40%, including infanticide. A significant positive correlation was found between the age of an infant at death or disappearance and the mother’s subsequent interbirth interval. Five cases of social change are described, including female transfer, one-male to multimale change, troop formation, and gradual and rapid replacement of troop males. Takeover-associated infant killing by band males, in an undisturbed moderate-density population, supported the sexual-selection/infanticide hypothesis but not the social-pathology hypothesis. However, it could not be directly confirmed that an invading infanticidal male gains a reproductive advantage. The male tenure of harems was estimated to be 45 months.     

Attacks on a wild infant ring-tailed lemur (Lemur catta) by immigrant males at Berenty, Madagascar: interpreting infanticide by males

An orphaned infant ring-tailed lemur (Lemur catta) was attacked persistently by immigrant males and disappeared with severe wounds at Berenty Reserve, Madagascar. Prior to the attacks, two troop members disappeared. On 20 November 1998, the only resident male in Troop C2B disappeared suddenly. After the disappearance, nine males from three neighboring troops approached, but only six males continued to shadow the troop. Of the females, the one lactating female attacked the males the most frequently. On 21 January 1999, the lactating female disappeared and her infant was orphaned. Subsequently, five attacks on the infant by immigrant males were observed for five consecutive days from January 23 to 27. The aggression was persistent and specifically targeted the infant, suggesting that the attacks were purposeful aggression, rather than redirected or accidental aggression. The primary attacker was the most dominant of the immigrant males, and mated with females in the next mating season.     

Take-over and infanticide in South Indian Hanuman langurs (Presbytis entellus)

A take-over and infanticide were observed in a South Indian population of Hanuman langurs (Presbytis entellus). A one-male, bisexual troop of langurs (Troop B1) was attacked by a band of two adult males and one subadult male. During the take-over, the resident male and one elderly female were driven from the troop. The youngest infant, about two months old, was fatally wounded by an attack from the dominant incoming male (S1). However, other young infants in Troop B1 were left unharmed. All three incoming males remained in Troop B1 but the troop appeared to remain functionally one-male, as S1 did not allow the other adult male to copulate. There was no sign that S1 attempted to eject the other two males from troop B1. © 1993 Wiley-Liss, Inc.