Mothers matter! Maternal support,dominance status and mating success in male bonobos (Pan paniscus)

Variation in male mating success is often related to rank differences. Males who are unable to monopolize oestrous females alone may engage in coalitions, thus enhancing their mating success. While studies on chimpanzees and dolphins suggest that coalitions are independent of kinship, information from female philopatric species shows the importance of kin support, especially from mothers, on the reproductive success of females. Therefore, one might expect a similar effect on sons in male philopatric species. We evaluate mating success determinants in male bonobos using data from nine male individuals from a wild population. Results reveal a steep, linear male dominance hierarchy and a positive correlation between dominance status and mating success. In addition to rank, the presence of mothers enhances the mating success of sons and reduces the proportion of matings by the highest ranking male. Mothers and sons have high association rates and mothers provide agonistic aid to sons in conflicts with other males. As bonobos are male-philopatric and adult females occupy high dominance status, maternal support extends into adulthood and females have the leverage to intervene in male conflicts. The absence of female support to unrelated males suggests that mothers gain indirect fitness benefits by supporting their sons.     

Which adult male savanna baboons form coalitions?

We investigated the mechanism of alliance formation among adult male savanna baboons by comparing the characteristics of males that formed coalitions frequently with males that never or seldom took part in coalitions. We observed three groups: two of Papio cynocephalus cynocephalusin Amboseli National Park, Kenya, and one of P. c. anubisin the vicinity of Gilgil, Kenya. We considered four hypotheses: (1) Males must be familiar with each other, (2) males must have an affinitive bond, (3) males must have more than average experience, and (4) the combined fighting ability of the coalition partners relative to the fighting ability of their opponent determines the likelihood that a coalition is formed. We conclude that relative fighting ability forms the key factor in coalition formation. High- ranking males do not form coalitions often, since they hardly need them. Low- ranking males rarefy form coalitions, since they cannot form effective coalitions among themselves. Affinity (“friendship”) may play a role as an additional factor. The relation of coalition formation with age and period of residence, which was found in several studies, can be explained largely by the correlation between these parameters and fighting ability.     

Aggressive interventions and matrilineal dominance relations in semifree-ranging barbary macaques (Macaca sylvanus)

Female Barbary macaques (Macaca sylvanus) form matrilineal hierarchies, i.e. they come to rank below their mother in relation to non-kin females in the course of maturation. The stability of such hierarchies and the acquisition of the matrilineal rank are achieved through dyadic aggressions and third party interventions in conflicts. This study examines the dynamics of interventions in non-kin conflicts in a semifree-ranging group of 109 Barbary macaques at “La Montagne des Singes,” Kintzheim, France. Focal sampling on 13 females aged 3 and 4 years not yet dominant over all older females from lower ranking kin groups (lower born females) was carried out during 16 months in 1987 and 1988. Results on the direction of support in non-kin female conflicts (all samples pooled) show that interventions were generally provided on behalf of the female from the higher ranking kin group (higher born female). Rates of interventions (derived from focal samples) given and received were correlated with the hierarchy; higher born females received more support and intervened more often than lower born females. A subset of interventions (based on the age of the females involved) was analyzed according to the rank distance between the opponents and the type of support provided (spontaneous or solicited). On the basis of their representation, intermediate-ranking supporters (i.e. females ranking between the opponents) intervened more often than above-ranking supporters (i.e. females dominant to both opponents), and they intervened more often spontaneously than following a solicitation. The results on interventions are discussed in the perspective of benefits to supporters. Twenty-one instances of outranking of older females (matrilineal rank acquisition) were observed. By the end of the study, the number of older lower born females not yet outranked by the focal females was negatively correlated to the rank distance between the two sets of females. However no such correlation was found between these two groups when compared according to their age difference in years.     

Social structures in Pan paniscus: testing the female bonding hypothesis

Based on previous research in captivity, bonobos, Pan paniscus, have been called a female-bonded species. However, genetic and behavioural data indicate that wild females migrate. Bonding between these unrelated females would then be in contradiction with socio-ecological models. It has been argued that female bonding has been overemphasized in captive bonobos. We examine patterns of proximity, grooming and support behaviour in six well established captive groups of bonobos. We find that female bonding was not a typical characteristic of all captive bonobo groups. In only two groups there was a trend for females to prefer proximity with other females over association with males. We found no evidence that following or grooming between females was more frequent than between males and unrelated females or between males. Only in coalitions, females supported each other more than male–female or male–male dyads. We also investigated five mother–son pairs. Grooming was more frequent among mothers and sons than in any other dyad, but sons did not groom their mothers more than males groomed unrelated females. Mothers groomed their sons, or provided more support to them than females groomed or supported unrelated males. Thus, while bonds between females were clearly present, intersexual relations between males and either unrelated females or their mothers are of more, or equal importance.     

Female-female social relationships in wild white-faced capuchin monkeys,Cebus capucinus

A single social group of wild white-faced capuchin monkeys was studied for a period of 26 months at Lomas Barbudal Biological Reserve, Costa Rica. A total of 604 hr of focal animal data was collected on six adult females in a group of 21 monkeys. Females could be ranked in a stable, linear dominance hierarchy. Adult females spent much more time in proximity to other adult females than to adult males. Females groomed other females twice as often as they groomed males, and about 55 times more often than males groomed males. Females tended to groom up the dominance hierarchy, and dyads with smaller rank distances groomed more often. Higher-ranking females nursed infants other than their own at lower rates than did lower-ranking females; however, females nursed infants of females ranked both above and below them. Although lower-ranking females were more likely than higher-ranking females to be the victims of aggression, higher-ranking females were not necessarily more aggressive than lower-ranking females. In 96% of female-female coalitions vs. a female, the victim was lower-ranking than both coalition partners; in the remaining 4%, the victim was intermediate in rank between the two coalition partners. Higher-ranking female-female dyads formed coalitions more often than did lower-ranking dyads. Those female-female dyads that groomed more frequently also formed coalitions more frequently. The patterning of social interactions indicates that Cebus capucinus at Lomas Barbudal are female bonded. © 1996 Wiley-Liss, Inc.     

Agonistic Behaviour and Sexual Conflict in Atypical Reproductive Groups: The Case of Bearded Vulture Gypaetus barbatus Polyandrous Trios

Social behaviour in species forced to form atypical breeding coalitions is poorly documented. The saturation of optimum territories in the Bearded vulture Gypaetus barbatus population in the Pyrenees has led floating males to settle in already occupied territories, thereby forming polyandrous trios. We examined the patterns of intrasexual aggression in five trios (nine reproductive events in total) during courtship. Alpha males initiated 82% of agonistic encounters that were mainly aimed at preventing or disrupting copulations. During the fertile period in recently formed groups, intrasexual aggression had a negative influence on the frequency of heterosexual copulations, which may be a contributing factor to the lower productivity of polyandrous trios. Females rejected a higher proportion of beta male- than alpha male-initiated copulations, and rejected copulations with both alpha and beta males more often when the other male was present close by. These results indicate that alpha males cannot effectively prevent all copulation attempts by beta males and that females avoid harassment by minimizing sexual activity when both males are present. Aggression between males decreased with time, occurring less often in established than in recent trios, despite the fact that the frequency of heterosexual copulations – the cause of conflicts – was similar. The frequency of homosexual interactions tended to increase in established trios, suggesting that this behaviour may help to regulate aggression within these groups, although no significant relationship between homosexual interactions and aggression was found. In summary, reproductive conflicts in trios seem to be unavoidable, although they tend to decrease if the group is maintained. This suggests that, for birds in these groups, the maintenance of a quality territory is more important than solving sexual conflicts.     

Coalition formation,mate selection and pairing behaviour of the Crested Francolin

Transect field observations were conducted on the behaviour of Crested Francolin Dendroperdix sephaena to describe male coalitions in the Borakalalo National Park, North West province, South Africa during May, August, October and December 2008, and again in July 2009. Crested Francolin form male coalitions following the breeding season in May. Young males leave their natal coveys and join old males in these coalitions, which are not based on kinship and consist of up to nine members. These coalitions are joined by a young female from June to October, followed by the breeding season when pairs replace coalitions. The coalition is a prelude to successful mating. The single female calls in the coalition, which triggers a cacophony, and her presence often erupts in physical conflicts between males. This cacophony is interpreted as signal jamming where males prevent one another from forming a duet with the female, clearly showing competition between them for winning over the female for mating. A well-synchronised duet forms the basis of pair formation. Old females are not particularly selected for pairing. The coalition is also a surrogate covey for young males because it optimises predator surveillance. The pair bond also optimises predator surveillance for each other throughout the year.     

Gonadal development and an indication of functional protogyny in the Indian damselfish (Dascyllus carneus)

The objective of this study was to determine the sexual pattern of the Indian dascyllus Dascyllus carneus . After an initially undifferentiated state, gonads of D. carneus developed an ovarian lumen and primary growth stage oocytes, and subsequently cortical-alveolus stage oocytes. From ovaries with cortical-alveolus stage oocytes and from more developed ovaries, some gonads redifferentiated into testes. From a sample of 163 individuals, two had a gonad containing degenerating vitellogenic oocytes and proliferating spermatogenic tissue, nine had a gonad containing degenerating cortical-alveolus stage oocytes and spermatogenic tissue, and five had a gonad with degenerating primary growth stage oocytes and spermatogenic tissue. The size of these individuals overlapped greatly with the size range of mature females, suggesting that at least in some individuals, redifferentiation toward a testis occurred after spawning as females. This indicates that D. carneus is a functional, diandric protogynous hermaphrodite. Removal of a dominant male(s) did not induce a sex change in any of the ranking females in the laboratory and field groups. There was no difference in the number of chases and signal jumps performed by the ranking female between control and experimental field groups, or before and after removal of the male. However, the sizes of the ranking females were at or beyond the size range of individuals with a mixed-stage gonad, suggesting that the developmental window for female-to-male sex change may not be open ended. In 41 of 43 field groups, in which sex of fish was determined histologically or by the shape of the urogenital papilla, one to several highest size ranks were occupied by males, followed by one to several females. Mature males, however, were not limited to the highest ranks and occurred at various lower size ranks within groups. Individuals with a mixed-stage gonad also occupied various size ranks within groups.     

Which adult male savanna baboons form coalitions?

We investigated the mechanism of alliance formation among adult male savanna baboons by comparing the characteristics of males that formed coalitions frequently with males that never or seldom took part in coalitions. We observed three groups: two of Papio cynocephalus cynocephalusin Amboseli National Park, Kenya, and one of P. c. anubisin the vicinity of Gilgil, Kenya. We considered four hypotheses: (1) Males must be familiar with each other, (2) males must have an affinitive bond, (3) males must have more than average experience, and (4) the combined fighting ability of the coalition partners relative to the fighting ability of their opponent determines the likelihood that a coalition is formed. We conclude that relative fighting ability forms the key factor in coalition formation. High- ranking males do not form coalitions often, since they hardly need them. Low- ranking males rarefy form coalitions, since they cannot form effective coalitions among themselves. Affinity (“friendship”) may play a role as an additional factor. The relation of coalition formation with age and period of residence, which was found in several studies, can be explained largely by the correlation between these parameters and fighting ability.     

Dominance status of adult male Japanese macaques: Relationship to female dominance status,male mating behaviour,seasonal changes,and developmental changes

Aggressive dominance orders of all adults in a confined troop of Japanese macaques (Macaca fuscata) were determined each mating and birth season during a 4-year interval. Males outranked more females in the mating than in the birth season, and some males shifted back and forth from ranks lower than female ranks in the birth season to ranks higher than female ranks in the mating season. Mating behaviour (number of female partners in mount and ejaculation series and ejaculation frequency) did not differ among males with ranks higher than, as high as, or lower than those of most females, nor did individual males mate more in years, when they were high-ranking than in years when they were not. There was a correlation, however, between ejaculation frequency and the number of females defeated by males. A pattern of increasing male rank with age was found.     

Social correlates of testosterone and ornamentation in male mandrills

We investigated relationships between fecal androgen concentrations, facial coloration and behaviour in semi-free-ranging male mandrills. We found that fecal androgen levels were significantly positively related to dominance rank, independent of rank stability and the mating period, suggesting that male mandrills live in a permanently aggressive context in which they must actively maintain their dominance status. Facial red coloration was also significantly related to both fecal androgen levels and rank, with high ranking males having both higher androgen levels and redder faces, although dominant males did not always have the highest androgen levels or the reddest faces. Predictive relationships between androgen levels, coloration and rank were short-term. Androgen concentrations and facial redness both increased in the presence of receptive females, as did the former during periods of rank instability. We conclude that male facial redness is likely to represent an honest signal (to other males) of current androgen status, competitive ability and willingness to engage in fights and that females may also use this to assess male condition. Further, our findings provide support for the “challenge hypothesis” as originally proposed for birds by Wingfield.     

Patterns of interventions and the effect of coalitions and sociality on male fitness

In group living animals, especially among primates, there is consistent evidence that high-ranking males gain a higher reproductive output than low-ranking males. Primate studies have shown that male coalitions and sociality can impact male fitness; however, it remains unclear whether males could potentially increase their fitness by preferentially supporting and socializing with females. Here we investigate patterns of male interventions and the effect of coalitions and sociality on male fitness in rhesus macaques (Macaca mulatta) with particular focus on male-female interactions. We combined behavioural observations collected on Cayo Santiago with genetic data analysed for male reproductive output and relatedness. Our results revealed that the ten top-ranking males provided the majority of all male support observed. In contrast to other primates, male rhesus macaques mainly formed all-down coalitions suggesting that coalitions are less likely used to enhance male dominance. Males supporting females during and before their likely conception were not more likely to fertilize those females. We also found no evidence that males preferably support their offspring or other close kin. Interestingly, the most important predictor of male support was sociality, since opponents sharing a higher sociality index with a given male were more likely to be supported. Furthermore, a high sociality index of a given male-female dyad resulted in a higher probability of paternity. Overall, our results strengthen the evidence that sociality affects fitness in male primates, but also suggest that in species in which males queue for dominance, it is less likely that males derive fitness benefits from coalitions.     

Agonistic aiding: Kinship,rank, age,and sex influences

An analysis of 3,774 episodes of agonistic aiding collected during a two-year study of a rhesus monkey group (Macaca, mulatta) indicated the differential influence of kinship and rank relationships on the participation of different age-sex classes in both aid to victims and aid to aggressors. Most aiding favored victims rather than aggressors and was much more likely to occur when matrilineal kin were involved. Females were more likely to aid than were males, and the frequency of their participation increased with age. Females were much more influenced by kinship than were males and defended or aggressively supported kin against any third party regardless of dominance relationships. Adult males seldom aided against animals that were dominant to themselves; the rare exceptions occurred when adult males defended kin. Aiding was far more likely to occur if the victim was squealing, and noisy agonistic episodes often involved multiple aiders on both sides. Aiding patterns had some potential to insure dominance rank inheritance within families, in accordance with the Kawamura hypothesis. In aiding animals outside of their own matrilines, however, group members aided randomly with respect to this model. There was little evidence that aiding functioned to support individuals when they targeted animals to which they should be dominant as adults based on matrilineal dominance relationships. Most defensive aiding seemed to function primarily to defend victims (primarily kin) of aggression. Aggressive support of the attacker, on the other hand, seemed to function primarily to reinforce coalitions with the attacker. The identity of the victim was unimportant as long as it was neither kin to nor dominant to the aider. Aggressive support of attackers did not overturn existing dominance relationships.     

Reproductive effort of bison bulls (<Emphasis Type="Italic">Bison bison</Emphasis>) in semi-natural conditions

Reproductive effort (RE) is defined as the investment in any current reproductive act. In male ungulates, variation in RE can be explained by differences in age, dominance, phase in the breeding season and number of oestrus females simultaneously present. We investigated the relation between these factors and RE in American bison males in semi-natural conditions. We looked for differences between males in RE, measured by tending behaviour, and maintenance activities (feeding/resting) during the rut. We found no effect of age on RE. During early rut there was no difference in RE between males of different rank. Dominant males had a higher RE during peak rut, and in late rut both the first and second ranking males showed more RE than lower ranking bulls. For all males, there was a clear increase in RE towards peak rut and when more females were attractive. Maintenance behaviours decreased when more females were synchronously in oestrus. The most dominant bulls spent less time grazing than lower ranking males, and all males decreased feeding during peak rut. Resting decreased with age. We conclude that dominance, phase of rut and the number of females synchronously in oestrus are the factors that best explain variation in RE.     

Sex, Rank and Age Differences in the Japanese Macaque (Macaca fuscata yakui) Participation in Inter-Group Encounters

In many species interactions among group are often characterized by agonistic behaviour. Although animals may participate in inter‐group encounters in different ways, depending on their energetic requirements, reproductive tactics, and/or developmental stage, the proximate causes affecting an animal's participation in inter‐group encounters are still poorly understood. Indeed, many studies have analysed the behaviour of males and females during inter‐group encounters without considering the importance of additional factors (e.g. rank). This study focuses on wild non‐provisioned Japanese macaques (Macaca fuscata yakui) living on Yakushima Island, Japan. It aims to determine how monkeys of different sex, age, and rank behave during inter‐group encounters and it discusses the implications and consequences of their behaviour on group composition and male dispersal. Males participated significantly more than females in inter‐group encounters, by displaying more aggressive or affiliative behaviour. High‐ranking and/or adult males were more aggressive than low‐ranking and/or subadult males during encounters occurring in the mating season and they also showed more herding behaviour. This trend was not found in inter‐group encounters occurring during the non‐mating season. Finally, males which then emigrated to new groups were low‐ranking and/or subadult individuals. Those males displayed more affiliative behaviour towards foreign males than males which did emigrate. These data indicate that in non‐territorial species with male dominance over female and high competition for mating partners males play an active, and often aggressive, role during inter‐group encounter while female participation is scarce. Factors such as age, rank and period of the year (in seasonally breeding species) have to be taken into considerations when analysing interactions between groups and their effects on group composition and social behaviour.     

Measuring male-female relationships during the mating season in wild Japanese macaques (Macaca fuscata yakui)

Heterosexual relationships during one mating season were examined in a wild troop of Japanese macaques (Macaca fuscata yakui) on Yakushima Island, Japan. Validation tests of putative mate choice behaviors demonstrated that female initiation and maintenance of proximity, female lookback at the male, and sexual presents to the male, were associated with increased mating. Male grooming the female was also associated with increased mating. Ten dyadic social behaviors were subject to principal components analysis to empirically define behavioral dimensions of male-female relationships. The analysis yielded four relationship dimensions: ‘Mutual Choice and Male Coercion,’ ‘Female Choice’ (two types), and ‘Mutual Choice’ Dyads tended to be characterized by more than one dimension. The results suggested that females sought matings with multiple males of various dominance ranks. Female relationships with high ranking males contained elements of male coercion and mate guarding, however, because these males attempted to inhibit females from mating with lower ranking males. The correlation between each relationship dimension and mating success depended, in part, on the dominance rank of males. Relationships involving high ranking males, which were most likely to contain elements of male coercion and mate guarding, were associated with mating success. Relationships involving low ranking males, which usually lacked such coercive elements. were less strongly correlated with mating success. These results, obtained from a wild troop, are compared to those previously obtained in captive and provisioned groups of Japanese macaques.     

Mechanisms of sexual selection: sexual swellings and estrogen concentrations as fertility indicators and cues for male consort decisions in wild baboons

Male mate-guarding episodes ('consortships'), are taxonomically widespread, yet costly to individual males. Consequently, males should bias consortships toward females with whom the probability of conception is high. We combined data on consortships with visual scoring of sexual swellings and assays of fecal estrogen concentrations (fE) in a wild population of baboons (Papio cynocephalus) to test the hypotheses that sexual swellings are reliable indicators of (1) within-cycle timing of ovulation, (2) differences in conception probability among females that differ in maturational stage, and (3) conceptive versus non-conceptive cycles of parous females. We also evaluated whether adult males might rely on swellings or other estrogen-dependent signals (e.g., fE) for mate-guarding decisions. We found that sexual swellings reflected conception probability within and among cycles. Adult males limited their consortships to the turgescent phase of cycles, and consorted more with adult females than with newly cycling adolescents. The highest ranking (alpha) males discriminated more than did males of other ranks; they (1) limited their consortships to the 5-day peri-ovulatory period, (2) consorted more with adult than with adolescent females, and (3) consorted more with adult females on conceptive cycles than on non-conceptive cycles, all to a greater extent than did males of other ranks. Male mate choice based on sexual swellings and other estrogenic cues of fertility may result in sexual selection on these female traits and enhance dominance-based reproductive skew in males. Alpha males are the least constrained in their mating behavior and can best take advantage of these cues to mate selectively.     

Male collared lizards,Crotaphytus collaris (Sauria: Crotaphytidae), signal females by broadcasting visual displays

The signalling function of displays broadcast when animals are distant from conspecifics can be difficult to determine. I tested the extent to which visually transmitted broadcast displays given by free‐ranging territorial male collared lizards signalled same‐sex rivals or females. One test involved recording the frequency of broadcast displays, aggressive contests with rivals, and courtship encounters with females during ten reproductive seasons when local sex ratios varied markedly. The frequency of broadcast displays decreased as the ratio of male competitors to females increased. The frequency with which males initiated contests with rivals was not related to the ratio of competitors to females, whereas the frequency of courtship interactions decreased with sex ratio because there were fewer females to court. The behaviour of males that defended territories during two successive seasons showed a similar pattern. Broadcast display frequency was positively correlated with courtship frequency, but not with the frequency of contests with rivals. Lastly, individual males gave more broadcast displays during focal observations when they also engaged in courtship encounters with females than other observations when they engaged in aggressive conflicts with rival males. Although these results do not reject the possibility that broadcast displays may also signal male rivals, they support a major role of these displays in advertisement to females. © 2013 The Linnean Society of London     

Aggression and worker control of caste fate in a multiple-queen wasp, Parachartergus colobopterus

Although famously cooperative, social insect colonies harbour considerable potential for genetic conflict among colonymates. This conflict may be expressed behaviourally as aggression by workers. We investigated aggression in 34 colonies of the wasp Parachartergus colobopterus, by evaluating the characteristics of both instigators and victims of aggressive interactions. We estimated genetic relatedness and queen number using DNA microsatellites and found that workers and emerging females should be most in conflict over the caste of the latter when there are many queens on the nest. We found that aggressive interactions are more likely to involve older workers attacking either males or younger workers, and that victim and aggressor females have more ovarian development than randomly sampled colonymates. Moreover, mated females with low levels of ovarian development relative to active queens were also more likely to be aggressors and victims than were randomly sampled females. Aggression among females supports the hypothesis that older workers use aggression towards younger females as a means of policing the development of emerging females into queens. Workers also may use aggression to suppress the reproduction of some mated females. Our findings thus support the hypothesis that genetic conflicts of interest motivate worker aggression in swarm-founding wasp colonies.