Natal emigration by both sexes in the La Pacifica population of mantled howlers: when do some stay?

We have reported previously that all male and female mantled howlers emigrate from natal groups at Hacienda La Pacifica, Costa Rica. In the years since that report, a small number of juveniles have stayed in the natal group without experiencing a solitary phase. Here, we present a post hoc analysis on juvenile emigration in six groups of howlers under observation for varying amounts of time between 1972 and 2005. Our records revealed 139 juveniles for whom emigration status was certain, and 125 of these did emigrate. There was a significant association between presence of mother and emigration: juveniles without mothers were more likely to remain in their natal group (chi(1)(2) = 53.1, P<.0001). The mean age of emigration for all juveniles (n = 125) was 2.47 years (SD = 0.9, range = 1.5-6.5). There was no difference in age of emigration by adult male composition (one-male, multi-male, both), but juveniles of unknown sex emigrated younger than either known males or females (F(2,116) = 4.4, P<.02). For emigrating juveniles of known sex (n = 99), both males and females without mothers left at a later age than those with mothers (F(1,95) = 6.5, P<.02). Although philopatry or delayed emigration occurs in a few motherless animals, most males and females do emigrate from their natal groups at ages consistent with those reported for other species of howlers.     

Dispersal Patterns Among Olive Colobus in Taï National Park

In Primates, females are more likely to be philopatric than males. However, in some species like Procolobus verus, females or individuals of both sexes disperse. In Taï National Park, Ivory Coast, olive colobus groups are small, with one or two adult males and 6 females. Dispersal is common for juveniles and adults of both sexes. Adult male dispersal is less common than adult female dispersal. Adult females immigrated especially into small, one-male groups indicating that food competition played a role. Furthermore, unknown sexually receptive females visited resident groups and mated with the resident males for a few days before disappearing again. Adult males dispersed when this improved their mating opportunities. All juveniles left their natal groups. The dispersal of juveniles may be a strategy to prevent inbreeding with their parents. Dispersal by juvenile males furthermore seemed to be the result of mate competition. The high dispersal rates, visits by receptive females, and dispersal of all individuals in the population suggest that moving between groups is a strategy that can be used ad hoc in several situations more easily in the olive colobus than in most other primates. The predation risks related to moving between groups were reduced by dispersing in conspecific or allospecific groups and by dispersing to neighboring groups.     

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.     

Male Migration in Barbary Macaques (Macaca sylvanus) at Affenberg Salem

We analyzed male migration during a 20-year period in the free-ranging Barbary macaque population of Affenberg Salem. Most natal migrations occurred around puberty, but only one third of all males left the natal group. Secondary group transfers were rare. All males immediately transferred to other bisexual groups. Migration rates were highest during periods with high adult female/male ratios within social groups. Immigrants highly preferred groups with fewer males of their own age than in the natal group, and many males immigrated into groups that had no male their own age. These groups originated from a skewed distribution of resident males during group fissions. A comparison of emigrants with their natal peers supports the inbreeding avoidance hypothesis as cause of emigration rather than the male competition avoidance hypothesis. Emigrants had no lower individual rank position and did not come from lower-ranking matrilines. Emigrants had more female maternal relatives, especially sisters. Males without female relatives almost never emigrated. Conversely, there is virtually no indication that emigrants were evicted from the natal group. Emigrants had no increased mortality. Paternity data revealed that the reproductive success of emigrants and natal males is similar, indicating that emigration had no reproductive cost. Many similarities between emigrants and natal males that separated from female maternal kin during group fissions suggest that inner migration during fissions is an alternative way to avoid maternal inbreeding. The mating system resulted in a genetic structure within social groups that largely diminished the chances for paternal inbreeding even without recognizing paternal kin.     

Group Fission in Barbary Macaques (Macaca sylvanus) at Affenberg Salem

We analyzed eight group fissions occurring during a 20-year period in three groups of a free-ranging provisioned Barbary macaque population. The founder group fissioned four times within 3.5 years after transfer to the enclosure, indicating that external factors—new environment, more space, absence of other groups—facilitated group fissions. Two groups resulting from these fissions, split twice within 2.5 and 1 years, respectively, many years later. The process of fissioning lasted from a few months to almost 2 years. Fissions were preceded by peripheralization/subgrouping of mainly young adult males (8-10 years old), suggesting that male competition was the primary force for the fissions. The males were joined by middle- to low-ranking but not the lowest-ranking females. The resulting new groups were usually smaller than the groups in which the former -matriline—old groups—stayed, and they were also more variable in size and sex ratio, suggesting that variable numbers of surplus individuals were expelled during fission. Mean adult sex ratios were similar in both groups after fission, indicating that the competitively superior males in the old groups (groups + -matriline) could not increase their breeding opportunities. Female kin, even of large matrilinies, almost always stayed together during fission. Natal males strongly preferred to join the old groups, and this preference was most pronounced in juveniles and subadults. Hence, most natal males stayed with maternally related females, i.e., remained true natal males, if the females stayed in old groups. They were separated from female kin, i.e., became seminatal, if the females joined the new groups. These seminatal males did not differ from natal males with respect to matrilineal rank, but they had more female relatives, above all more close relatives (sisters), indicating that avoidance of mating with maternal kin was important for group choice. Despite joining the same group as female kin during fissioning, breeding opportunities of natal males (ratio of unrelated females/male) were not less than that of their seminatal peers, because natal males had fewer female relatives. Only a minority of both groups of males would have done better by joining the alternative group. Paternal relatives were distributed during fission by chance, and loss of patrilinies was therefore much less pronounced. We conclude that the rules governing social relationships among Barbary macaque males are less apt to cope with the high number of males resulting from provisioning, whereas the rules regulating social relationships of females living in a nepotistic, female-bonded society are very robust in this respect.     

Effects of dispersal status on pituitary and gonadal function in the male spotted hyena

Male spotted hyenas (Crocuta crocuta) reach puberty at 24 months of age and then usually emigrate from their natal clans one to 52 months later. Recent work has shown that reproductive success is very low among adult males still residing in their natal groups, and it is similarly low among recent or "short-term" immigrants. Long-term immigrants father the vast majority of cubs born. Here we inquired whether these differences in reproductive success might be associated with variation among males in pituitary or gonadal function. In one free-living hyena population in Kenya, we compared baseline levels of plasma luteinizing hormone (LH) and testosterone (T) among adult natal males, short-term immigrants, and long-term immigrants, and we also compared their pituitary and gonadal responses to GnRH challenge. Mean basal plasma LH values did not differ among groups, but mean basal T concentrations in long-term immigrants were higher than those in either natal males or short-term immigrants. Similarly, pituitary response to GnRH challenge did not vary significantly among groups, but testicular response to challenge was greater in long-term immigrants than in natal males or short-term immigrants. Thus adult natal and immigrant males exhibited similar pituitary function but testicular function was attenuated among adult males that had not yet left their natal groups and also among males attempting to become established in new social groups. This suggests that, like the act of emigration itself, the process of social integration during immigration may have important physiological consequences for male mammals transferring between groups.     

Behavior and endocrine concentrations do not distinguish sex in monomorphic juvenile howlers (Alouatta palliata)

Behavioral observations on juvenile mantled howlers are limited by visually undifferentiated genitalia; however, animals can be sexed when they are very young or if they are captured. Behavioral data and fecal samples from juveniles during month-long field studies from 1993 to 1995 were analyzed to determine whether there are developmental differences in behavior or hormone concentrations that can be used to differentiate males from females. The subjects were juveniles of known sex and age from five different social groups on Hacienda La Pacifica, Costa Rica. Based on 749.8 hr of focal-animal sampling, there were no sex differences in daily activity patterns. There were no sex differences in proximity to mothers and other group members, and age differences reflected howler life-history patterns. There were no differences in estradiol or testosterone concentration by age or sex. Juvenile monomorphy thus extends beyond morphology to behavioral and hormonal similarity as well. Most juveniles are forced out of their natal groups and remain solitary until they join new groups by supplanting all same-sex adult group members. Monomorphy may allow them to spend more time in natal groups, and thus both reduce the solitary period and allow the juveniles to improve social skills needed for later immigration. While this strategy may benefit juvenile howlers, it remains a problem for those who wish to study juvenile sex differences from a distance.     

Origin and fate of a one-male savanna babbon group formed by fissioning

An adult male chacma baboon,Papio hamadryas ursinus, emigrated from his habituated natal group at age 100 months and joined a neighboring multimale-multifemale group. Subsequently this male, together with eight adult females, all with infants <1 year old, formed a one-male group (OMU). The male was the only adult male in this group for over 200 days. Circumstantial evidence shows that shared parentage of the infants with this male determined which females joined the new group. In addition, 10 juveniles younger than 4.5 years joined the new group. Based upon persistent close physical association, especially at sleeping sites, these juveniles were presumed to be the older independent progeny of the eight adult females. Thus, the founding elements of the new group were the adult male and partial matrilines—mothers, their infants, and their independent juvenile offspring. The OMU status of the group ended with the sequential intrusion of three males known to be low ranking in other groups. The first of these intruders was successful in joining the OMU in spite of the OMU male's earlier successes in rebuffing this and other potentially high-ranking males. Six additional males later entered the fission group during a 6-month interval. Two of them outranked the original male. All additional males present in the last 100 days of observations came from the same group, which was not the group of origin of the females.     

Socioecological factors of male chimpanzee migration at Bossou, Guinea

For more than 21 years a small semi-isolated group of wild chimpanzees have been studied at Bossou, Guinea, west Africa. All individuals have been identified since the beginning of the study. Remaining rates of infants (0–3 yr) and juveniles (4–7 yr) in the group were 64–80% for both sexes, however, those of adolescents (8–11 yr) dropped drastically, particularly for males (14%). As a result, most natal males as well as females disappeared before fully maturing. Two male visitors and an immigrant were observed in the group. More adult males than females disappeared from Bossou. Group males could be excluded as the genetic father of an offspring born in the group. From these demographic trends it is highly likely that some of these males emigrated rather than succumbed to sickness and death. It seems likely that they left on their own by choice. The reason for male dispersal is hypothesized to be influenced by intra-group male-male competition and the habitat ecology and structure of Bossou. There are no competitive adjacent groups or predators to prevent males from living alone and males can sire offspring out of their natal group.     

Sons of low-ranking female rhesus macaques can attain high dominance rank in their natal groups

Five adult and subadult sons of middle- and low-ranking female rhesus macaques (Macaca mulatta) were observed to hold high dominance rank in their natal groups during a 12-month study at Cayo Santiago, Puerto Rico. Three of these males also experienced high mating success during at least one mating season. These findings contrast with all previously published accounts of rank acquisition by natal male rhesus macaques in provisioned colonies, and they present a challenge to the hypothesis that natal transfer functions to increase male access to fertile females.     

无量山大寨子黑长臂猿一夫二妻制的群体结构及其行为学原因

2003年8月—2005年8月,对无量山大寨子5个黑长臂猿群体的结构和组成进行了观察。当一个群体在早晨鸣叫或依次通过树冠时,记录群体的结构和组成。每个群体都由1个成年雄性、2个成年雌性及其后代组成。2003年8月平均群体大小为6·2只;到2005年8月,平均群体大小发展为6·4只,其中有2个亚成年雄性从出生群迁出,且有3只幼猿出生。在3个群体(G1、G2和G3)中两个成年雌性都成功繁殖了后代。同一群体内两个成年雌性间无攻击或等级行为。2005年4月15日,当一只亚成年雌性进入G3的领域后,两只成年雌性对其进行追逐驱赶,并且干扰其与成年雄性配合进行二重唱,成年雄性没有直接驱赶流浪的亚成年雌性,10天后这只亚成年雌性离开了G3的领域。亚成年雄性经常与群体其他成员保持一定距离,并且在出生地通过独唱练习鸣叫。黑长臂猿可能通过亚成年雄性和雌性的迁出,及成年雌性对外来流浪雌性的驱赶维持这种一夫二妻的群体结构。     

Does intraspecific competition facilitate age separation in timing of southward migration in waders?

In many Palaearctic wader species there is a clear separation in the timing of adult and juvenile southward migration. This phenomenon is traditionally explained by the selection on adults to depart early from breeding grounds and necessity of juveniles to prepare longer for migration. In this study we hypothesize that late departure from natal grounds may also be adaptive for juveniles, as it allows them to avoid intensified interference competition at stopover sites with adult, usually more dominant conspecifics. To test this hypothesis we analysed long-term data on stopover behaviour of juvenile wood sandpipers (Tringa glareola) staying at a central Polish stopover site under varying levels of competition from adult birds. The results clearly indicated that juveniles were highly disadvantaged by the simultaneous presence of adults at the same staging site, as under intense competition from older conspecifics they refuelled more slowly and attained lower fat reserves. It was also found that juveniles which were forced to compete with adults left the site quickly and possibly searched for more favourable staging places. All these imply that delayed departure from natal grounds may be adaptive for juvenile waders, allowing them to mismatch the timing of their first migration with the peak of adult passage and, thus, reduce the negative consequences of intraspecific competition during migration.     

Facultative dispersal by juvenile males in the cooperative stripe-backed wren

We present genetic and demographic data documenting juveniledispersal in the cooperatively breeding stripe-backed wren (Campylorhynchusnuchalis) of Venezuela. Parentage and DNA fragment-sharing analysesrevealed 12 cases in which juveniles were unrelated to othergroup members. Of these 12 foreign juveniles, (1) all were males,(2) eight of 12 had been found with breeding pairs lacking helpersrather than with groups containing helpers, and (3) four outof seven of those observed as adults courted or sired offspringwith the dominant females in their new groups despite the strongincest avoidance of this species. Furthermore, juvenile maleshad a significant tendency to disappear from natal groups intheir first year, and singleton juveniles observed with pairsafter the breeding season were mostly males. These data supportthe hypothesis that foreign juveniles were dispersers from intactgroups and not products of conspecific brood parasitism or adoptionfollowing group dissolution. We suggest that unassisted pairsmight accept juvenile males into their groups as helpers toincrease their future reproductive success and that dispersersthemselves might leave large natal groups in which their helpingis superfluous to join small groups of nonrelatives in whichthey might soon reproduce.     

Juvenile dispersal in Calomys venustus (Muridae: Sigmodontinae)

Both spacing behaviour and dispersal movement are viewed as hierarchical processes in which the effects may be expressed at spatial scale. This research was carried out to examine the hypothesis that the presence of parents promotes the dispersal of juveniles from their natal nest and their father or mother home-range, in Calomys venustus.The study was carried out in four 0.25 ha fences (two controls and two experimentals), in a natural pasture. This study had two periods: Father Removal (FR) (August and December 1997; year one) and Mother Removal (MR) (August 1998 and January 1999; year two). For the FR treatment fathers were removed after juveniles were born, but in the MR treatment mothers were removed after the juveniles were weaned. The effect of parents on the dispersal distance of juveniles was analysed with respect to their natal nest and their mother and father home-range. Dispersal distance from the nest of C. venustus was independent of either male or female parent. Juveniles were more dispersing in relation to the centre of activity of their mothers than to that of their fathers, and females were more dispersing than males. Female juveniles overlap their home-range with their parents less than male juveniles do. The differences observed between female and male juveniles would be related to their different sexual maturation times, as well as to the female territoriality.     

Male life history and intergroup mobility among ringtailed lemurs (Lemur catta)

Lemur catta shares a male-biased dispersal pattern with most primate species and the majority of mammals. Individuals in a free-ranging population of ringtailed lemurs were captured, marked, released, and monitored for a 40-month period. Sixty-four percent of the males (43 of 67) migrated or were missing within this period and all nine censused groups were affected by migration. Males migrate from their natal group and then may migrate again after reaching full adult body weight. Full-sized adult males migrate at a rate of 0.28 per year or once every 3.5 years and may change groups a number of times during their life. Migrations occurred within a 6-month period, ending just after the mating season. However, there is no direct connection between mating success and male migration. Females mate with transferring males, with group males, and with visitors from adjacent groups. The age-related pattern of male migration and the occurrence of extragroup mating inLemur catta is similar to that described for several species of macaques.     

Life history of hamadryas baboons: Physical development,infant mortality,reproductive parameters and family relationships

Demographic and life history parameters were estimated for a band of free-ranging hamadryas baboons, observed for 5.5 years in Ethiopia. Age-related changes in body weight and dentition were found to be delayed relative to laboratory-reared baboons. On the average, females reached menarche at 4.3 years of age and had their first infant at the age of 6.1 years. The mean interbirth interval was 24 months if the infant survived this period. The survival of infants and juveniles was higher compared to Amboseli yellow baboons, but somewhat lower than in gelada baboons in the Simen Mountains. Males acquired their first juvenile or adult female at the age of 8.5 to 11 years. Male-female pair-bonds lasted several years in most cases. The Cone Rock baboons were organized in a four-level social structure. The troop could split into bands, bands were divided into clans, and clans into one-male units with bachelor followers. The exchange of individuals between social units predominantly occurred within the band. All males of known origin became adult members of their presumed natal clan. Most females transferred also within clans, and juvenile females tended to remain in their natal clan. Females lost by one male to several rivals tended to reassemble in the same new one-male units later on.     

Geographic variations in underwater male Weddell seal Trills suggest breeding area fidelity

Adult Weddell seals (Leptonychotes weddellii) exhibit site fidelity to where they first breed but juveniles, and perhaps transient adult males, may disperse from their natal location. If there is mixing between adjacent breeding groups, we would expect that common vocalizations would exhibit clinal patterns. Underwater Trill vocalizations of male Weddell seals at Mawson, Davis, Casey, McMurdo Sound, Neumayer and Drescher Inlet separated by ca. 500 to >9,000 km, were examined for evidence of clinal variation. Trills are only emitted by males and have a known territorial defense function. Trills from Davis and Mawson, ca. 630 km apart, were distinct from each other and exhibited the greatest number of unique frequency contour patterns. The acoustic features (duration, waveform, frequency contour) of Trills from Neumayer and Drescher Inlet, ca. 500 km apart, were more distinct from each other than they were from the other four locations. General Discriminant Analysis and Classification Tree Analysis correctly classified 65.8 and 76.9% of the Trills to the correct location. The classification errors assigned more locations to sites >630 km away than to nearest neighbours. Weddell seal Trills exhibit geographic variation but there is no evidence of a clinal pattern. This suggests that males remain close to single breeding areas throughout their lifetime.     

Group takeover by a natal male howling monkey (Alouatta palliata) and associated disappearance and injuries of immatures

As part of a long-term study on howling monkey behavior and social dynamics, a known natal male was observed taking over his group from his putative sire. Due to the accidental death of one of the adult males, this natal male had matured in a one-male group and had never observed juvenile male emigration nor adult male immigration and associated behaviors. Nevertheless, the behaviors associated with the takeover were indistinguishable from those of an immigrant male, including disappearance of immatures, one of whom was found with extensive injuries. While it cannot be said that the natal male inherited these behaviors from his presumed father, it can be said that he exhibited species-typical behaviors associated with male takeover in the absence of observational learning.     

Adult-young interactions in island and mainland populations of the deermouse Peromyscus maniculatus

Summary The demographic and ecological characteristics of island populations of small mammals have received increasing attention in recent years, but few studies have compared the behavioral characteristics of island populations with those of mainland populations. Behavior is considered an important variable because it is believed by many to be a crucial factor affecting the population dynamics and demography of natural populations. In particular, among many species of rodents, the social behavior of adults towards juveniles is cited as an important factor influencing dispersal patterns and population regulation. The present study compares social interactions between adults and juveniles of island and mainland populations of the deermouse Peromyscus maniculatus, and attempts to relate differences in behavior to the demographic differences between the two populations. Adult mice were trapped on the mainland of British Columbia and on one of the Gulf Islands off the British Columbia coast, and allowed to breed in the laboratory. Male and female juveniles from both populations were then tested with their own parents and with unrelated male and female adults. The results demonstrate that island adults show almost no aggression towards either own or unrelated young. Mainland adults likewise show little aggression towards their own young, but a proportion of the population, consisting of both male and female adults, shows severe aggression towards unrelated juveniles of both sexes. These results suggest four major conclusions: 1) behavior may be the mechanism responsible for the demographic differences reported for these island and mainland populations; 2) female aggression may be a more important factor in deermouse population dynamics than has been previously recognized; 3) since parents show little aggression towards their own young, adult aggression may be a significant factor in juvenile mortality and emigration only after juveniles have initiated dispersal away from their natal sites; and 4) adult aggression controls the number of both male and female juveniles which are recruited into the population.     

Behavioral and social correlates of escape from suppression of ovulation in female common marmosets housed with the natal family

Although female common marmosets typically do not breed while housed with their natal families, up to half ovulate at least once while housed with the intact natal family, and a similar proportion conceive if an unrelated adult male is present in the group. In this study, we investigated the behavioral and social correlates of escape from suppression of ovulation by daughters housed in intact natal families or in families in which the father had been replaced by an unrelated adult male. Focal-animal behavioral data were collected from daughters that were (N = 7) or were not (N = 10) undergoing ovulatory cycles while housed with the natal family and from daughters that were (N = 5) or were not (N = 3) cycling or pregnant in families containing an unrelated male. Additionally, four cyclic and six acyclic females housed in intact natal families underwent simulated “prospecting” tests. Cyclic and acyclic daughters in intact natal families did not engage in sexual interactions with the father and showed few differences from one another in their interactions with the parents. Moreover, cyclic and acyclic daughters did not differ in their willingness to leave the family for short periods or to investigate an unfamiliar family in “prospecting” tests. However, daughters that underwent ovarian cycles in the presence of an unrelated male showed numerous behavioral differences from those in intact natal families, including frequent courtship and sexual behaviors with the male, reduced affiliative interactions with the mother, and elevated frequencies of aggressive display behavior. Moreover, these females were less likely to behave submissively towards the mother or the adult male. These findings suggest that both suppression of ovulation and inhibition of sexual behavior normally contribute to reproductive failure in female marmosets living with their natal families, and that the two components of suppression may become dissociated under specific social conditions. Am J Primatol 41:1–21, 1997. © 1997 Wiley-Liss, Inc.