You are Not Welcome: Social Exchanges between Female Spider Monkeys (<Emphasis Type="Italic">Ateles geoffroyi</Emphasis>)

Group living leads to competition for food between group members. Two types of intragroup food competition may occur: scramble competition, in which all group members use the same resource, such that feeding opportunities are equal for everyone; and contest competition, in which some group members monopolize resources through aggression and dominance. In species in which females disperse from the natal group and immigrate into other groups, immigrant females increase group size and thus possibly food competition. Under these circumstances, other females may use aggression to discourage new females from joining the group. We assessed the distribution of aggression, embraces, and kisses among female spider monkeys (Ateles geoffroyi) in relation to group tenure. We recorded social interactions during 1688 10-min focal animal samples on 11 females in Santa Rosa, Costa Rica. We found that aggression was rare between long-term resident females and aggression rates were not higher during feeding than in other contexts, suggesting there was little contest competition. Long-term residents and less recently immigrant females showed higher aggression rates toward the most recent immigrants than toward other females, especially during the first months after a female immigrated, which coincided with the dry season. We did not find similar patterns for embrace and kiss. These results suggest that other females target aggression toward the most recent immigrants to reduce scramble competition. This finding suggests that group tenure should be included in socioecological models for species with female dispersal.     

A comparison of the mating behavior of adolescent and adult female rhesus macaques (Macaca mulatta)

This study compares adult and adolescent female rhesus macaques with regard to (1) characteristics of their copulatory partners, (2) their proceptive behaviors, and (3) adult male behaviors toward them during estrus. We conducted focal follows of 24 adolescent and 65 adult free-ranging estrous female rhesus macaques on Cayo Santiago during two mating seasons. Compared to adult females, adolescents presented sexually to males at higher rates; copulated more frequently with rankless young male, and extra-group males; and, in one of two mating seasons, were ignored more frequently by males to whom they presented sexually. Adolescents tended to copulate with ranked, resident males at higher frequencies on days when the operational sex ratio (adult males:estrous adult females) was high. Males directed “muzzle-up” signals to adolescents at lower rates than to adults in one of two mating seasons, although this effect vanished when males who might have fathered adolescent females were excluded from analysis. Adolescents did not differ consistently from adults in strength of the correlation between proximity maintenance (dyadic Hinde's Index) and copulation rate, or in approach rate to males. Adolescent females, relative to adult females, presented sexually more to rankless young males, but did not present more to ranked, resident males. Both proximate (e.g. endocrine) and ultimate (e.g. differential fecundity; female-female mate competition) explanations may account for the reported differences between adult and adolescent female rhesus macaque sexuality.     

Dynamics of fission in a wild Barbary macaque group (Macaca sylvanus)

We studied the dynamics of a group of Barbary macaques (Macaca sylvanus) in Algeria from March 1983 to November 1989. Troop fission began in autumn 1987, when group size had more than doubled, to include 76 animals. We observed 11 temporary splits of this group during the mating seasons of 1987 and 1988. The process was interrupted during the 1988 birth season. In June 1989, fission resumed and ended with the formation of three independent groups that included 50, 24, and 13 individuals. Adult females played an important role in the process of fission. They initiated the rapid formation of two, and later three, coherent nuclei, distributed in two or three bisexual subgroups; on several occasions these nuclei also formed subgroups without any adult males. Adult males remained together in a single nucleus for longer periods of time than females did. However, during fission, 35% (N=20) of resident males emigrated to neighboring groups, while 11 strange males immigrated into the focal group; over 6 years, 57% of the male transfers occurred after the beginning of the process. After group fission, maternally related individuals lived together in the new groups. The majority of resident males remained with the largest of the three groups, while most of the immigrant males were in another group. The third group included a single adult male. Possible factors that induce group fission are discussed.     

Social relationships between immigrant and resident bonobo (Pan paniscus) females at Wamba.

Social relations and behaviours of adolescent female pygmy chimpanzees (Pan paniscus) that migrated between unit groups were studied at Wamba, Za?re. Each immigrant female selected one particular resident female, a 'specific senior female' (SSF) and frequently approached and followed her. Affinitive behaviours (e.g. grooming, genitogenital rubbing) were frequently performed between the immigrant and her SSF. Most affinitive behaviours were more likely to be initiated by the immigrant than by her SSF. The immigrant's relationship with her SSF seemed to facilitate the immigrant's integration into the new unit group. Resident males actively approached and mated with the immigrant females. Copulation also appeared to further the immigrant's stable relationship with the resident males.     

Troop Extinction and Female Fusion in Wild Japanese Macaques in Yakushima

We observed three cases of troop extinction and two cases of female fusion in the wild population of Japanese macaques on Yakushima Island, Japan. Troops P and T decreased in size relatively slowly over a few years until each troop consisted of only three monkeys. Several months later, the remaining adult female of P merged with the adjacent troop S, followed by the remaining female of T. S subsequently also decreased in size and disappeared about 2 years later. In the early stage of troop decline, the mortality rate of adult females was as low as in a growing troop, but the birthrate was quite low. In the late stage of troop decline, the mortality rate increased and the birthrate remained low. An important factor leading to troop extinction may be an increase in population density and the resulting increase in intergroup competition. During the period when P and T declined and ceased to exist, the range of the adopted troop shifted to cover their previous ranges. In the fused troop, there was no severe aggression directed towards the immigrant females or harassment from residents of the adopted troop and there was affiliative social interaction between the immigrant females and resident members. These results agree with previous reports on female fusion: it occurs when the shrinking group consists of one or no adult member, and the immigrant females are not at a severe disadvantage in their adoptive group. A possible benefit for immigrant females is to avoid disadvantage of one-adult group in conflict with conspecifics. A possible cost for immigrant females is transfer to the other troop or to unfamiliar area or both. The cost to transfer to another group may not be high because the members of the adoptive troop are relatively tolerant to immigrants. The cost to transfer to unfamiliar range may be minimized by immigration to the troop whose range shifted to the immigrants' former range.     

Non-reproductive copulation behavior among Tibetan macaques (<Emphasis Type="Italic">Macaca thibetana</Emphasis>) at Huangshan,China

Non-reproductive copulation, which takes place outside of the mating season and does not result in conception and birth, was studied in a free-ranging group of Tibetan monkeys (Macaca thibetana) at Mt. Huangshan, China, in the birth seasons of 1992 and 1997. We employed all occurrence and focal animal samplings to record sexual and related behaviors and affiliation interactions, respectively. Compared with sexual behavior in mating season, non-reproductive copulation occurred at a lower frequency, with less frequent ejaculation, less harassment, shorter mount duration, and an absence of pause with vocalization. It often took place in a situation in which non-lactating females were involved in social conflict or approached males for mating. Neither pregnant nor lactating females were observed to mate in the birth seasons. Copulation during the birth season did not increase a sexually receptive female’s delivery the next year, nor was it associated with increased proximity, grooming, or agonistic aid for the mating pair. However, copulated pairs spent more time co-feeding, presumably reflecting an increased tolerance on the part of the male. Adolescent males, who rarely copulated in the mating season, engaged in mating activity in birth seasons as well. Therefore, though birth-season copulation had no reproductive functions, it may have fulfilled social functions for females, such as post-aggression appeasing by males or gaining access to resources. This also offered good opportunities for adolescent males and females to develop their sexual skills for later competition.     

Kin recognition in a social spider

Social spiders accept immigrant spiders into their kin-based groups, suggesting that spiders cannot recognise kin and may lose inclusive fitness benefits. A field and two laboratory experiments on Diaea ergandros, a social crab spider, demonstrated that younger and older instar D. ergandros do discriminate siblings, but potential benefits were variable and not equally distributed. First, proportional survival was greater in large groups regardless of the within-group relatedness, so accepting immigrants increases probability of group survival (although relatedness was more important among smaller groups). Second, juvenile D. ergandros ate unrelated spiders instead of siblings when starved, so immigrants might represent a food reserve in times of food shortage. Third, subadult resident, sibling females cannibalised unrelated, immigrant females and their brothers instead of immigrant males when starved, suggesting that subadult female spiders may maximise outbreeding opportunities. These benefits provide selective pressure for groups to accept immigrants, but as benefits are realised differentially, conflict and cooperation will exist within spider groups similar to that shown in other group-living taxa.     

Proximal Spacing Between Individuals in a Group of Woolly Monkeys (Lagothrix lagotricha) in Tinigua National Park, Colombia

I studied proximal spacing within a group of woolly monkeys (Lagothrix lagotricha) during 7 months at Parque Nacional Tinigua, Colombia. I collected a total of 1188 instantaneous samples on focal individuals, recording the number and age/sex class of individuals that were in contact with, <2 m from, <5 m from the focal animal. The results indicate that proximate spacing reflects social affinities and is related to mother–infant relationship and social grooming. Subadult females and adult males are the sex/age classes with the lowest number of individuals in proximity. There are low proximity between adult females and between adult males and high frequencies of nearness between mother and offspring. Associations between males and females were usually low, but in some cases males showed preferences for a given female. There was a relatively gradual increase in spacing between mothers and their offspring as they became older. Old juvenile males were associated chiefly with other males—mostly subadults—whereas juvenile females maintained some proximity only to their mothers. There are also differences in spacing behavior according to different activity types.     

Dispersal patterns in Costa Rican mantled howling monkeys

Both male and female juveniles disperse in Costa Rican mantled howling monkeys (Alouatta palliata). 79% of the males and 96% of the females leave their natal groups. Males may spend up to 4 years and females up to 1 year as solitaries. Extra-group individuals are faced with only three possibilities, i.e., form a new group by joining another extra-group individual, join an established social group, or remain solitary. Most surviving extra-group individuals join an established social group which contains no kin. Females join with the help of a resident male and once in a group proceed to rise to the alpha position through dyadic interactions. The immigrant female either becomes the alpha female or leaves and tries again in another group. Males challenge the alpha male and either defeat him or remain solitary. Competition with relatives for limited high quality food may be the reason for both sexes leaving their natal groups in howlers. By leaving, the successful immigrants increase their mothers inclusive fitness while suppressing the fitness of nonrelatives instead of remaining natal and competing with relatives for limited food.     

Gene flow and immigration rate in an island population of great tits

It is generally recognized that immigration and gene flow cannot be equated, but few detailed quantitative comparisons of these processes have been made over the entire lifetime of individual animals. We analyzed data collected in a longterm study of an island population of great tits Parus major, and tested two assumptions frequently made in population genetic studies. (1) The assumption that there is no difference in reproductive output between immigrant and resident breeding birds was refuted for females but not for males. Lifetime production of local recruits (LRS) was reduced by 37% in immigrant females, because female immigrants tended to leave the island after breeding, and thus reproduced for fewer years. Female LRS was density dependent, but the effect of density was independent of status (resident/immigrant). (2) The assumption that mating was random with respect to status was also refuted: assortative mating was found, even when temporal and spatial aggregation of immigrants was controlled for. Thus both assumptions were refuted, and gene flow was lower than immigration rate.     

The fitness consequences of kin-biased dispersal in a cooperatively breeding bird

Cooperative alliances among kin may not only lead to indirect fitness benefits for group-living species, but can also provide direct benefits through access to mates or higher social rank. However, the immigrant sex in most species loses any potential benefits of living with kin unless immigrants disperse together or recruit relatives into the group in subsequent years. To look for evidence of small subgroups of related immigrants within social groups (kin substructure), we used microsatellites to assess relatedness between immigrant females of the cooperatively breeding superb starling, Lamprotornis superbus. We determined how timing of immigration led to kin subgroup formation and if being part of one influenced female fitness. Although mean relatedness in groups was higher for males than females, 26% of immigrant females were part of a kin subgroup with a sister. These immigrant sibships formed through kin recruitment across years more often than through coalitions immigrating together in the same year. Furthermore, females were more likely to breed when part of a kin subgroup than when alone, suggesting that female siblings form alliances that may positively influence their fitness. Ultimately, kin substructure should be considered when determining the role of relatedness in the evolution of animal societies.     

Seasonal, sex, and interspecific differences in activity time budgets and diets of patas monkeys (Erythrocebus patas) and tantalus monkeys (Cercopithecus aethiops tantalus), living sympatrically in northern Cameroon

I examined seasonal, sex, and interspecific differences in activity time budgest and diets of patas (Erythrocebus patas) and sympatric tantalus monkeys (Cercopithecus aethiops tantalus) on the basis of 5-day data sets collected in three and two different seasons, respectively, by the method of focal animal sampling. The seasons included species-specific mating and birth seasons. As compared with not only the birth season but also conspecific females, both patas resident male and tantalus male spent less time feeding and more time resting, day and night, in their respective mating seasons. Given that day-resting time includes time for vigilance for non-resident males and receptive females, this may reflect that males should minimize time spent feeding to allow maximum participation in other fitness-increasing activities such as mating-relating activities asSchoener (1971) predicted. In both species, the males consumed fruits containing less protein but more calories and showed a high feeding rate to compensate for the shorter time spent feeding in the mating season. In contrast, females consumed protein-rich food types (i.e. animals, protein-rich seeds, leaves, and flowers) in the birth season to meet the high demand for protein due to pregnancy and lactation. Given that the season for males was considered to be not a calendar but a reproductive “season” (i.e. mating or birth season), both sexes of patas spent more time moving and less time day- and night-resting than did the tantalus counterparts irrespective of the “season”. Patas subsisted on fruits, gums, and supplementarily lipid-rich seeds as an energy source and animal matters and protein-rich seeds as a protein source. In contrast, tantalus subsisted on fruits and lipid-rich seeds as energy and flowers and leaves as protein.     

Severe Aggression Among Female Pan troglodytes schweinfurthii at Gombe National Park, Tanzania

Aggression is generally more severe between males than between females because males gain greater payoffs from escalated aggression. Males that successfully defeat rivals may greatly increase their access to fertile females. Because female reproductive success depends on long-term access to resources, competition between females is often sustained but low key because no single interaction leads to a high payoff. Nonetheless, escalated aggression can sometimes immediately improve a female’s reproductive success. Resisting new immigrants can reduce feeding competition, and infanticide of other females’ young can increase a female’s access to resources. East African chimpanzees live in fission-fusion communities in which females occupy overlapping core areas. Growing evidence indicates that reproductive success correlates with core area quality, and that females compete for long-term access to core areas. Here we document 5 new cases of severe female aggression in the context of such competition: 2 attacks by resident females on an immigrant female, a probable intracommunity infanticide, and 2 attacks on a female and her successive newborn infants by females whose core areas overlapped hers. The cases provide further evidence that females are occasionally as aggressive as males. Factors influencing the likelihood and severity of such attacks include rank and size differences and the presence of dependable allies. Counterstrategies to the threat of female aggression include withdrawing from others around the time of parturition and seeking male protection. We also discuss an unusual case of a female taking the newborn infant of another, possibly to protect it from a potentially infanticidal female.     

Resistance Evolution to Bt Crops: Predispersal Mating of European Corn Borers

Over the past decade, the high-dose refuge (HDR) strategy, aimed at delaying the evolution of pest resistance to Bacillus thuringiensis (Bt) toxins produced by transgenic crops, became mandatory in the United States and is being discussed for Europe. However, precopulatory dispersal and the mating rate between resident and immigrant individuals, two features influencing the efficiency of this strategy, have seldom been quantified in pests targeted by these toxins. We combined mark-recapture and biogeochemical marking over three breeding seasons to quantify these features directly in natural populations of Ostrinia nubilalis, a major lepidopteran corn pest. At the local scale, resident females mated regardless of males having dispersed beforehand or not, as assumed in the HDR strategy. Accordingly, 0–67% of resident females mating before dispersal did so with resident males, this percentage depending on the local proportion of resident males (0% to 67.2%). However, resident males rarely mated with immigrant females (which mostly arrived mated), the fraction of females mating before dispersal was variable and sometimes substantial (4.8% to 56.8%), and there was no evidence for male premating dispersal being higher. Hence, O. nubilalis probably mates at a more restricted spatial scale than previously assumed, a feature that may decrease the efficiency of the HDR strategy under certain circumstances, depending for example on crop rotation practices.     

Group Composition Affects Seasonal Birth Timing in Captive Japanese Macaques

We analyzed birth dates recorded during an 18-year period in a group of Japanese macaques housed in the Rome zoo to assess the influence of environmental, physiological, and social factors on birth seasonality. Birth timing differed significantly among years. Birth timing was affected by reproductive condition of females—ones that had given birth in the previous year delivered significantly later than those that had not—but not by their age or dominance rank. We conducted further analyses separately on females that had or had not given birth in the previous year. In both subgroups of females mean birth date was not influenced either by environmental temperature and rainfall during the previous mating season or by group size. On the contrary, among females that had not given birth in the previous year, socionomic sex ratio—ratio of sexually mature males to sexually mature females—is positively correlated with both mean birth date and date of the first birth, but not with date of the last birth. Contrarily, among females that had given birth in the previous year, there is no significant relationship between these variables. We hypothesize that the effects of socionomic sex ratio on birth timing might depend on competition among males for access to fertile females. When the number of males per female was higher, mutual disruption of consort pairs may have led to a delay in the onset of mating.     

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.     

Spawning periodicity and annual breeding seasons of Tilapia nilotica (L.) in the White Nile

Two annual breeding seasons are indicated for populations of T. nilotica in the White Nile: a major autumnal season (mid-July–September) coinciding with the annual rainfall and a smaller back-up season in February–March. During these seasons reproducing females constituted 80% and 40% of the total population of mature females compared to 10–12% in the intervening summer period. Enhanced gonadal development, fecundity and GSI values characterized the breeding seasons. Gonadal maturation proceeded uninterruptedly even during the summer and repeated fractional spawning is suggested for this species within and beyond the breeding seasons. Most of the spring breeders were either very young or relatively old fish and most medium-sized females bred in the main autumnal season. Older fish were less fecund (by 70–75%) compared to younger fish. In the latter, fecundity increased progressively with body growth (r² = 0.732; p<0.001); the proportionality was lost in medium-sized fish and in older females (0.9–1.5 kg) fecundity correlated negatively with increased body size.     

Female social dynamics in a provisioned free-ranging band of the Sichuan snub-nosed monkey (Rhinopithecus roxellana) in the Qinling Mountains, China

Little information is available on the social dynamics of the Sichuan snub-nosed monkeys (Rhinopithecus roxellana). Here, we provide 5-year observational data on a provisioned free-ranging band in the Qinling Mountains, central China, from October 2001 to December 2005. The results of this study suggested that females place a high priority on maintaining long-term relationships with other female in their one-male units (OMUs). Females had more social interactions with unit members than with nonunit individuals. They showed relatively high rates of affiliation with their unit females, intermediate levels with the resident males, and low levels with individuals outside the unit. Females had few interactions with other resident males. Social interactions, especially affiliative interactions, were more symmetrical between focal females and their unit females than between focal females and the resident males. Thus female social interactions appeared to resemble those of other female-bonded primate societies. On the other hand, female transfers occurred sometimes. We recorded 11 cases of female immigration and 5 cases of female disappearance, 1 case of female intraband shift, and 2 cases of "merging of OMUs," which suggested that unit females might be loose-knit in this species. We discussed influencing factors on female Sichuan snub-nosed monkey social dynamics, by comparing them with those of Hamadryas and gelada baboons that also have multilevel societies.     

Female transfer between one-male groups of proboscis monkey (<Emphasis Type="Italic">Nasalis larvatus</Emphasis>)

Successful or unsuccessful female transfers were observed seven times during a 32-month field study of proboscis monkeys (Nasalis larvatus) inhabiting a riverine forest along a tributary of the Kinabatangan River, Sabah, Malaysia. In all cases, the females voluntarily left their own groups and immediately joined with another one. When adult females tried to shift to other groups, adult males called them back to their own groups, but appeared to be indifferent to subadult females. When the adult females returned, the males never attacked the females physically, but instead often emitted herding sounds to them. One subadult female was repelled by a resident adult female. When one adult female transferred into a new one-male group, she left her behind son in an all-male group. The number of females often fluctuated in most study groups, with this fluctuation being more prominent among subadult females than adult females. It is likely that female transfer in proboscis monkeys is not a rare occurrence and that it is especially common among sub-adult females.     

Breeding migrations by bighorn sheep males are driven by mating opportunities

In some species where male mating success largely depends on intrasexual competition, males can adopt migratory or resident strategies to seek breeding opportunities. The resulting mixture of resident and migrant tactics within a population can have important ecological, genetic, and evolutionary consequences for metapopulations. Bighorn sheep Ovis canadensis males establish a linear dominance hierarchy that influences their mating tactics. Some males perform breeding migrations during the pre‐rut and rut to seek mating opportunities, but little is known about these seasonal movements. We analyzed presence/absence data for 62 marked bighorn males during six mating seasons (20–32 males/year) in the Sheep River Provincial Park, Alberta, Canada, where hunting was not allowed. On average, about half of males left their natal population to rut elsewhere. The proportion of males leaving (yearly range 15%–69%) increased as the number of resident mature males increased and the populational sex ratio decreased, with fewer females during the pre‐rut. Among those leaving the park, 24% did so in October, while the trophy sheep hunting season was open. Detailed monitoring of breeding migrations in protected populations could inform management strategies to limit evolutionary impacts of hunting, which can alter size‐dependent mortality and create artificial pressures driving changes on heritable traits.