Alternative routes to the leader male role in a multi-level society: follower vs. solitary male strategies and outcomes in hamadryas baboons

The nested one-male units (OMUs) of the hamadryas baboon are part of a complex social system in which "leader" males achieve near exclusive mating access by forcibly herding females into permanent consortships. Within this multi-level social system (troops, bands, clans and OMUs) are two types of prereproductive males--the follower and solitary male--whose different trajectories converge on the leader role. Here we compare OMU formation strategies of followers, who associate with a particular OMU and may have social access to females, with those of solitary males, who move freely within the band and do not associate regularly with OMUs. Data were derived from 42 OMU formations (16 by followers and 26 by solitary males) occurring over 8 years in a hamadryas baboon band at the Filoha site in Ethiopia. "Initial units" (IUs) with sexually immature females (IU strategy) were formed by 44% of followers and 46% of solitary males. The remaining followers took over mature females when their leader was deposed (challenge strategy) or disappeared (opportunistic strategy), or via a seemingly peaceful transfer (inheritance strategy). Solitary males took over mature females from other clans and bands, but mainly from old, injured or vanished leaders within their clan (via both the challenge and opportunistic strategies). Former followers of an OMU were more successful at taking over females from those OMUs than any other category of male. Despite this advantage enjoyed by ex-follower leaders, ex-solitary leaders were equally capable of increasing their OMU size at a comparable rate in their first 2 years as a leader. These results demonstrate the potential for males to employ both multiple roles (follower vs. solitary male) and multiple routes (IU, inheritance, challenge, opportunistic) to acquire females and become a leader male in a mating system characterized by female defense polygyny in a competitive arena.     

Five natural troops of Japanese monkeys on Shodoshima Island: II. A comparison of social structure

The results of a comparative study of five natural troops of Japanese monkeys living on Shodoshima Island are as follows. (1) When troops were feeding at the feeding places, various cases, such as individuals spaced out dispersively or close by in a gregarious state, were observed. (2) Differences were also observed among the troops at the feeding place; some troops always fed in a dispersive state, on the whole, while others fed in a gregarious state with small distances between individuals. (3) Differences of social structure were observed among the five troops: differences of the number of classes and sub-classes in males and females, of the relation between classes and their age distribution in males, of the proportion of individuals who intruded into the central part of the troop or who made it their core area of activity to the total male population, etc., and it was also assumed that some troops were integrated more strictly, while others were integrated loosely. (4) These differences of social structure among the troops were assumed to be closely related to the gregariousness or dispersiveness of the each troop at feeding time. From these facts several decisive factors of social structure were considered.     

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.     

Population and social dynamics changes in ring-tailed lemur troops at Berenty,Madagascar between 1989 - 1999

In the present study, we recorded all births, immigrations, deaths, and emigrations for a population of ring-tailed lemurs at Berenty Reserve, Madagascar, between September 1989 and August 1999. In September 1989, three troops (C, B, and T) inhabited the study area of 14.2 ha. During the 10-year period, eight troop divisions, six evictions of females, and three troop takeovers of ranges by other troops occurred in and around the study area. Consequently, in August 1999, the number of troops in the same area increased to six (CX, C1, C2A, C2B, T1, and T2). The number of lemurs aged >1 year increased from 63 to 82, which resulted from 204 births, 58 immigrations, 125 deaths, and 118 emigrations. Of the 204 newborn lemurs during the study period, 103 died, 44 emigrated outside the study area, and 57 remained within the study area. The total number of lemurs that emigrated from natal troops was 69 (54 males and 15 females). Natal males left their troops around the age of 3. Non-natal males changed troops after a tenure varying from 1 to 7 years. Survival curves showed a fall in survival rates of both sexes to < 0.5 between the ages of 2 and 3. For females, the survival rate gradually decreased to < 0.2 at the age of 9. On the other hand, due to emigration, the survival rate of males could not be determined after the age of 5 yr. Since some males attained high-rank at the age of 6 – 10 yr, the prime age for male ring-tailed lemurs is thought to be around 7 – 10 yr. Ring-tailed lemurs are essentially female philopatric, because all cases of females leaving natal troops resulted from troop divisions or forced evictions. Such social changes may have resulted from competition among females. All cases of troop divisions or evictions occurred in larger troops consisting of ≥20 lemurs, and only a few females could rejoin their troops. When males joined such a female-group, a new troop was formed. Although promoted by an increase in population, frequent emigrations of females from original troops are the characteristics of ring-tailed lemurs at Berenty.     

An advantage for young sperm in the house cricket Acheta domesticus

We show that males of the house cricket Acheta domesticus regularly expel sperm packages (spermatophores) independently of copulation and at a rate that is not affected by the presence of females. We then show for the first time that the age of sperm affects their likelihood of being stored by females after copulation; younger sperm were overrepresented in the female sperm storage organ and therefore in the sperm population used for fertilization. Our results suggest that the reproductive success of males may increase if they deliver ejaculates with young sperm, and the results may explain why the males of several species are regularly observed to discard ejaculates. Our results also suggest that phenomena such as female multiple mating, paternity bias, and/or exaggerated ejaculate sizes may be related to the advantage both genders gain by using young sperm.     

Population organization of wild pig-tailed macaques (Macaca nemestrina nemestrina) in West Sumatra

A field study on wild pig-tailed macaques was conducted in West Sumatra, Indonesia, during three periods from January 1985 to February 1987. During the nine months of the first two periods, unprovisioned monkeys were traced and observed. During the eight months of the last period, monkeys were provisioned and observed mainly at baiting sites. Three troops and ten solitary males appeared at the two baiting sites. Some males immigrated into and emigrated from the troops. The troops had a multi-male multi-female composition. The size of the various troops was 74, 49, and 81 individuals, respectively, and the mean adult sex ratio in the troops was 1:6.3; that is, markedly biased towards females. The home ranges of two of the troops overlapped considerably. When the troops encountered each other at the baiting sites, a clear dominance relationship was recognized. The troops differed in their integration as ranging units: two of the troops did not form subgroups (temporary fission and fusion of each troop), while the other troop frequently split into subgroups. Recent field studies on pig-tailed macaques have suggested a multi-leveled society with harem-type unit groups. However, in the present study, the troops observed had neither a substructure similar to harem-type groups nor a superstructure that emerged as a result of fusion of the troops. The unit group of the pig-tailed macaques appears to be a multi-male, matrilineal group.     

Who is Responsible for Fission in a Free-ranging Troop of Baboons?

The study troop of chacma baboons (Papio cynocephalus ursinus) at Mkuzi Game Reserve, Zululand, South-Africa, comprised of about 76 members that split into two new troops. The events leading to this troop fission will be described and its possible causes will be discussed. Troop fission among baboons is generally attributed to the withdrawal of low-ranking females from the main group, as a result of the cost of food competition and its effect on their reproductive success. At Mkuzi, no evidence for food competition among females was recorded in terms of rank-related time spent feeding or other time—budget components, feeding-bout length, diet composition or context of female aggression. Moreover, no evidence for rank-related differential reproductive success was found in terms of inter-birth intervals or infant survival. Female mortality was, however, related to dominance rank, with circumstantial evidence suggesting that cause of mortality was predation by leopards. Rate of female disappearances, aggression levels among females, and the percentage of time they spent in proximity to other adult troop members increased after fission. Relatively short inter-birth intervals and extremely low infant mortality rate at Mkuzi resulted in a small number of receptive females at any one time, and therefore in high costs of male sexual competition as expressed in the high levels of male aggression and woundings, both reduced after fission. It is suggested that this troop fission may have been initiated by the resident males, triggered by the high cost of sexual competition, and forced on the females, who were, consequently, subjected to higher risk of predation. The troop fission was preceded by a long process of increasing tendency for sub-trooping. It was initiated by the four resident males who kept a large distance apart from each other, herded oestrous female associates away from others and were followed by other females. The females generally tended to stay close to associates, males and females. These parties were followed by the peripheral and immigrant males who had no female associates, and eventually two distinct daughter troops were formed.     

Sex differences in the use of daily torpor and foraging time by big brown bats (Eptesicus fuscus) during the reproductive season

Temperate zone bats can use daily torpor as a means of saving energy. Some argue, however, that torpor is costly for both males and females and that individuals should only use it during times of poor foraging conditions. Others hypothesize that the costs are greater for females and that males should enter torpor more regularly. We tested these alternative hypotheses by using temperature-sensitive radiotransmitters to monitor use of torpor and foraging by free-ranging big brown bats ( Eptesicus fuscus ). During the pregnancy period, males used torpor at night more and foraged less often than did females. Males also went into deep torpor more often and remained in torpor longer than did females. When they foraged, males and females were away from the roosts for equal periods of time. During the lactation period, males and females rarely failed to forage and foraging times were again no different between the sexes, although males may roost at night away from the maternity colonies. Males again used torpor and deep torpor more often and for longer than females did. These results support the hypothesis that the fitness costs of using torpor are lower for males than for reproductive females and that males regularly use torpor as an energy-saving mechanism. Females enter torpor only when foraging conditions are poor, presumably because torpor prolongs gestation and slows neonatal growth thereby leaving less time for females and their young to prepare for hibernation.     

RESOURCE INFLUENCED SEX ROLES OF ZAPROCHILINE TETTIGONIIDS (ORTHOPTERA: TETTIGONIIDAE)

Male tettigoniids donate nutrients to females at mating in the form of a spermatophylax. Male-donated nutrients function as paternal investment leading to a reversal in the sex roles of males and females. Reversal in the behavioral sex roles of a zaphrochiline tettigoniid was found to be directly related to the current availability of food resources in the environment. When resource availability was low, females were less fecund and males had lower and more variable accessory gland weights (the spermatophylax producing gland) than when resource availability was high. When resources were scarce, larger individuals had a reproductive advantage having more eggs or heavier accessory glands. All individuals were equally fecund or had equal accessory gland weights when resources were plentiful. During low resource availability males that had a spermatophylax to offer were choosy of their mates, and females were competitive. When resources were plentiful, males were less discriminative and females showed signs of discrimination. There was evidence that female sexual motivation decreased when environmental resources were plentiful (as indicated by mating frequency) and that only females of low nutritional status continued to mate. When resources were scarce, females achieved fecundities equivalent to those achieved during high resource availability through spermatophylax consumption. These data support the hypothesis that females continue to mate to compensate for low resource availability and that male parental investment may be important only in poor quality habitats. When resources are plentiful females do not need to remate. Thus resource availability may directly influence the number of sexually active males and females (operational sex ratio) and the form of sexual selection.     

The relation between male body size, fighting, and mating success in Dawson's burrowing bee, Amegilla dawsoni (Apidae, Apinae, Anthophorini)

Males of the bee Amegilla dawsoni (Anthophorini) vary greatly in size, with some of the largest individuals as large as the largest females, a rare phenomenon in insects. The occurrence of unusually large males appears to be the product of sexual selection for fighting ability. As predicted from this hypothesis: (1) males regularly competed aggressively to be in the best position to mount sexually receptive, virgin females as they emerged from the ground; (2) larger males usually won fights for potential mates, as demonstrated by the fact that males able to defend sites with an emerging female were larger on average than males they kept at bay; (3) the fighting advantage of large males translated into greater mating success. Males captured while mounted on a female were significantly larger on average than randomly captured matesearching males in two of three populations sampled in 1993. Moreover, males known to mate more than once were consistently larger than single-mating males in the 10 samples taken from four populations examined in 1994. Thus, a large male mating advantage applies across years and among populations, making it potentially advantageous for females of Dawson's burrowing bees to produce large, superior fighting sons about the same size as their daughters.     

Social interactions among adult male langurs (Presbytis entellus) at Rajaji Wildlife Sanctuary

A population of langurs (Presbytis entellus)at the Rajaji Wildlife Sanctuary in northern India was investigated for 1820 hr throughout a 10-month period in 1978. Data were collected from four bisexual troops and the adult males that ranged outside of bisexual troops. Most (60%) of the observation hours occurred with a main study troop from which social and ecological data were collected. The langur population at Rajaji shows pronounced birth and mating seasons. The population density is high (ca. 80/km ²), with about 75% of the adult males living outside of bisexual troops, which typically are large and multimale. Males outside of bisexual troops occur in small all-male bands or as isolates. Relations between bisexual troops and all-male bands are characterized by relatively low levels of aggression, and members of all-male bands are able to associate with bisexual troops for prolonged periods during the mating season. As a result of these associations, nontroop males are about as successful as troop males in achieving reproductive access to troop females. These associations between bisexual troops and all-male bands occurred with a minimal amount of agonistic behavior and without mortality or injury to troop females or immatures.     

Symmetrical patterns in non-agonistic social interactions found in unprovisioned Japanese macaques

Non-agonistic social interactions in an unprovisioned troop of Japanese macaques (Macaca fuscata yakui) were analyzed with the spacing between individuals, leading-following interactions, and exchange of social grooming. The most frequent interactions were found between kin-related females. Unrelated females stayed with one another rather frequently, but rarely exchanged social behaviors. Interactions between males and females were infrequent though they were occasionaly observed between high-ranking males and high-ranking females. Very frequent exchange of grooming was observed between males, and even high-ranking males exchanged grooming more frequently with males than with females. Most non-agonistic social interactions in the study troop were based on bidirectional exchange of social behaviors, in which no clear tendency relevant to dominance or sex was found; while in provisioned Japanese macaque troops, associations between males and females, between unrelated females, and between males were formed mainly be subordinates' active roles in associative behaviors. This seems relevant to the idea that dominance grealty influence social life in provisioned troops. The present study provides guidelines for interspecific comparison of social interaction patterns of macaque species.     

Breeding and movements of goshawks in boreal forests in Sweden

Goshawks were studied in their boreal forest breeding habitat in central Sweden from 1977 to 1982. Nests were regularly spaced, with a nearest neighbour distance of 6.3 km. corresponding to a density of about 3 pairs 100 km⁻². Of initiated breedings. 68% were successful. There was no difference in breeding performance between years. Average brood size was 2.7. Sex ratio among first captures of adults was even, but males were recaptured more often than females. Nearly all juveniles left the area in winter. Among radio-tagged adults 20% of the males, and 50% of the females moved out of the area. When moving, both adults and juveniles moved into agricultural areas, the juveniles generally covering longer distances. From autumn to spring, body condition of females improved, whereas that of the males deteriorated. It is argued that the males' stronger tendency to stay in winter is due to the advantage of possessing a territory in early spring. Females, on the other hand, can move to areas with as good food conditions as possible.     

Can punishment maintain sex?

Individuals who reproduce asexually have a two-fold advantage over their sexually-reproducing counterparts as they are able to reproduce twice as fast. Explaining why sexual reproduction is favoured over asexual reproduction therefore remains an important challenge in evolutionary biology. Various mechanisms involving resistance to parasites, adaptation to novel environments and helping to purge the genome of deleterious mutations have all been proposed as potential mechanisms which could promote the evolution of sex. A recent article has suggested that spiteful males may help to reduce the two-fold advantage of asexual females. Here I discuss this idea, and further ask whether punishment of asexual females by sexual females could be one way in which sexual reproduction could be maintained in groups of animals; in light of recent research on the repression of competition, it could be possible that asexual females which reproduce faster than their sexual counterparts will be punished for using group resources. It may therefore be possible that the behaviour of sexual individuals towards asexual females could have fitness consequences which could potentially reduce the two-fold advantage they gain from reproducing parthenogenetically.     

Consortship and Mating Success in Chacma Baboons (Papio cynocephalus ursinus)

Chacma baboons (Papio cynocephalus ursinus) show a lower consortship take‐over rate and longer consortship duration than the other savannah baboons ( Bulger 1993 ). It has been argued that researchers have focused on atypically small troops with few adult males, resulting in low competition for access to oestrous females. Consortship data from two mountain baboon troops containing seven and four males, respectively, were analysed to determine whether the troop with the greater number of males showed a weaker correlation between mating success and rank due to an expected higher consortship take‐over rate. No consort take‐overs were observed in either study troop and mating success in both troops was correlated strongly with male rank. The distribution of days spent in consortship amongst the males could be explained by the priority‐of‐access‐model. The degree of cycle overlap determined the number of males observed consorting oestrous females, whereas the number of males did not influence the relationship between rank and consorting activity.     

Movement patterns of yellow baboons (Papio cynocephalus): Sex differences in juvenile development toward adult patterns

Age-sex differences in the sociospatial positioning of progressing baboon troops have been postulated as an important, protective aspect of baboon adaptation to a terrestrial existence. The protection theory is an ultimate hypothesis which can be disproven by showing that the postulated age-sex positioning fails to occur. Alternatively, confidence in the protection theory can be increased if the postulated positioning persists across several troops despite significant opportunities for variation, and if progression order is shown to be linked to biological phenomena such as sex differences in physical maturation. Data from 73 individually identified, free-ranging juveniles of both sexes were compared with previous findings. Small juveniles of several troops were more centrally located than large juveniles. Large juveniles males were more toward the front than were large juvenile females. As male walking infants and juveniles aged, a change in positioning (i.e., location) occurred: males were located increasingly toward the front of progressions. Comparable data from females suggest that the adult female pattern had begun to form among large juveniles. Data are consistent with an ultimate protection theory.     

Infanticide and juvenilicide in Hanuman langurs (Presbytis entellus) around Jodhpur,India

Ten cases of infant killings and 2 cases of juvenile killings were observed in two troops of Hanuman langurs, (Presbytis entellus) around Jodhpur, India. Fatal attacks on infants and juveniles are classified into four categories. The process of infanticide was observed from start to end and is described in detail for 3 cases. The age of the victims ranged from 0.2 to 48 months. The interbirth interval among females whose infants were killed is significantly shorter compared to females whose infants survived. In ourt study, 7 cases support the reproductive advantage hypothesis, that infanticide is an adaptive behaviour to increase male reproductive success. The remaining 5 cases do not fit into the reproductive advantage hypothesis. In these cases, victims are over 8 months old, and as such their deaths could not shorten the interbirth interval. It appears that by killing older infants and juveniles the males obtain an advantage in resource competition for their offspring. An alternative is that new males chase or peripheralise the older infants and juveniles, which leads to 97% predominant uni-male troop structure in Jodhpur.     

Shoaling and mate choice of wild-type Tanichthys albonubes in the presence of the red fluorescent transgenic conspecifics

Shoaling and sexual behaviour of wild-type male and female white cloud mountain minnow Tanichthys albonubes were measured in the presence of the red fluorescent transgenic conspecifics under laboratory conditions. Wild-type female test fish showed no significant preference, whereas wild-type male test fish preferred to be near a shoal of red transgenic fish rather than wild-type fish. When placed in a potentially reproductive context, wild-type males had a higher competitive ability over transgenic males; wild-type females spent more time with wild-type males in visually mediated experiments, but wild-type males performed more courtship displays towards transgenic females. These results suggest that the red body colouration does not appear to disturb signal communication between wild-type and transgenic T. albonubes in shoaling behaviour; transgenic males have no mating advantage over wild-type males, but the red body colouration of transgenic females may affect mate choice of wild-type males.     

Social Relationships Among Ring-Tailed Lemurs (Lemur catta) in Two Free-Ranging Troops at Berenty Reserve, Madagascar

We observed two free-ranging troops of ring-tailed lemurs at the Berenty Reserve, Madagascar. Kinship affinities in these troops are known only for mothers and their offspring 4 years of age. We attempted to quantify social relationships. Almost all agonistic interactions were dyadic, and triadic agonistic interactions, such as alliances, were very rare. Dominance hierarchies in both sexes in the two troops were not linear. As in cercopithecine monkeys, mothers were dominant over their adult daughters. However, the daughters were not ranked immediately below their mothers. Close proximity and social grooming occurred more frequently between closely related females, such as mother–daughter and sister–sister dyads, than between unrelated females. Frequent-proximity relations also occurred between adult males that had emigrated from another troop and entered the present troop together, even though they did not rank closely to one another. Subordinates were likely to groom and to greet dominants more frequently than vice versa. During group encounters, particular females were involved in agonistic interactions with animals of other troops, regardless of dominance rank. Adult males, regardless of their dominance rank, but not adult females, constantly tried to drive solitary males away.     

Sex differences in food hoarding behaviour of Long Evans rats

Food hoarding was examined in male and female Long Evans rats. During a three-week period the animals were regularly submitted to the testing procedures, with food pellets in the home cage being either unrestricted (first week), or restricted (second and third week). When food pellets were present ad libitum, the hoarding scores were low, lower in females than in males. When food deprivation was imposed (second week), the animals responded by hoarding more pellets. This increase was more pronounced in males than in females. During the third week the hoarding scores became more or less stable, with males hoarding significantly more pellets than females.