Looking like mother makes mallard ducklings dominant over their siblings

Colour variation in time and space among animals may affect social relationships such as pairing and dominance interactions. For instance, some birds are naturally sensitive to leg colour, with some colours being more visible or attractive than others. The colour of the leg-rings used to mark birds may thus be related to behavioural and reproductive variables. Most studies have investigated this effect for adults during reproduction, but leg-ring colour may also affect the behaviour of young birds. We tested the potential effect of leg-ring colours on the within-brood dominance hierarchy of mallard (Anas platyrhynchos) ducklings while each brood formed a stable and exclusive family unit with its mother. Ducklings did not acquire a within-brood dominance rank according to the colour of their own ring. This result suggests that mallards may not have a sensory bias for a given colouration. However, ducklings wearing a ring of the same colour as one of the two rings of their mother were dominant over their siblings. We discuss the potential behavioural and methodological implications of this result.     

The ghost of social environments past: dominance relationships include current interactions and experience carried over from previous groups

Dominance hierarchies pervade animal societies. Within a static social environment, in which group size and composition are unchanged, an individual's hierarchy rank results from intrinsic (e.g. body size) and extrinsic (e.g. previous experiences) factors. Little is known, however, about how dominance relationships are formed and maintained when group size and composition are dynamic. Using a fusion-fission protocol, we fused groups of previously isolated shore crabs (Carcinus maenas) into larger groups, and then restored groups to their original size and composition. Pre-fusion hierarchies formed independently of individuals' sizes, and were maintained within a static group via winner/loser effects. Post-fusion hierarchies differed from pre-fusion ones; losing fights during fusion led to a decline in an individual's rank between pre- and post-fusion conditions, while spending time being aggressive during fusion led to an improvement in rank. In post-fusion tanks, larger individuals achieved better ranks than smaller individuals. In conclusion, dominance hierarchies in crabs represent a complex combination of intrinsic and extrinsic factors, in which experiences from previous groups can carry over to affect current competitive interactions.     

Social behavior of the emperor tamarin in captivity: Components of agonistic display and the agonistic network

Agonistic behavior was studied longitudinally for 16 months in an intact family groups of captive emperor tamarins (Saguinus imperator subgrisescens) using methods from quantitative ethology and social network analysis. A motivational analysis of the components of agonistic display revealed the relative strength of each component along a continuum from strongly dominant to strongly subordinate. Tabulations of exchanges of strongly dominant and strongly subordinate components in interactions among the tamarins revealed an agonistic network (“dominance hierarchy”) that approached, but did not quite reach, the ideal state of a transitive order (“linear dominance hierarchy”). The frequency with which individual tamarins long called and scent marked was not closely correlated with their position (“dominance rank”) in the agonistic network. Instead, individuals undergoing change in status long called and scent marked frequently, irrespective of their rank.     

Agonistic dominance in male baboons: An alternative view

Data on baboons have generated both the concepts normally associated with male dominance hierarchies among primates and the tests of their significance. The priority-of-access model has been used to predict the relationship between dominance rank and resource acquisition. While the correlation between these two factors, or between rank and measures of reproductive success, has varied among different primate species, most recent baboon field-workers have interpreted their results to be consistent with the model. Based on 1200 hr of observation of a troop of savannah baboons near Gilgil, Kenya, this paper presents data on male agonistic interactions and on male acquisition of resources. Predictions of the priority-of-access model are tested and an inverse relationship is found between agonistic dominance rank and acquisition of two limited resources, estrous females and meat. The importance of the residency status of males is explored and an alternative hypothesis is presented to account for the anomalous pattern in the data. The relationship of male reproductive success and dominance rank is evaluated in light of the data on these baboons and the “residency” hypothesis.     

Testosterone and Linear Social Dominance Status in Captive Male Dabbling Ducks in Winter

Dominance hierarchies play an important role in avoidance and/or solving conflicts in gregarious species. In dabbling ducks (Anas species), dominance allows for feeding‐site monopolization in winter quarters where resources are generally limited. In addition, male social rank should theoretically favour access to mates. Dominance rank can be associated with morphological traits, and is often correlated with aggressiveness, a behavioural trait generally related to high testosterone levels. In this study, we investigated the existence of a winter group structure based on dominance relationships and tested for a linear hierarchy, in three species of captive male dabbling ducks (mallard Anas platyrhynchos, pintail A. acuta and wigeon A. penelope). We then analysed the relationship between dominance ranks, morphological parameters and testosterone levels measured in early (Oct.) and mid‐winter (Dec./Jan.). We found that the three male groups of the three species exhibited a linear hierarchy. Testosterone levels differed during winter and between species. Morphologic measurements, body mass and body condition were not correlated with individual dominance ranks, whereas dominant males had higher testosterone levels than subordinates. The slopes of the relationships were similar between species and winter period, but the y‐intercepts differed between species and between early and mid‐winter phases. The linear hierarchy found in the three species indicates that dominance relationships strongly structure dabbling duck groups in winter. Lack of correlation between rank and morphological characters, but correlation of rank with testosterone levels suggests that social rank is more dependent on behavioural traits such as aggressive behaviour. The differences between species and winter periods are discussed in relation to migration and wintering phenology.     

Survival of Wood Duck Ducklings and Broods in Mississippi and Alabama

ABSTRACT Although North American wood ducks (Aix sponsa) are well-studied throughout their range, researchers know little about demographic and environmental factors influencing survival of ducklings and broods, which is necessary information for population management. We studied radiomarked female and duckling wood ducks that used nest boxes and palustrine wetlands at Noxubee National Wildlife Refuge (NNWR) in Mississippi, USA, in 1996–1999, and riverine wetlands of the Tennessee-Tombigbee Rivers and Waterway (TTRW) system in Alabama in 1998–1999. We estimated survival of ducklings and broods and evaluated potentially important predictors of duckling survival, including age and body mass of brood-rearing females, hatch date of ducklings, duckling mass, brood size at nest departure, inter-day travel distance by ducklings, site and habitat use, and daily minimum air temperature and precipitation. At NNWR, survival of 300 radiomarked ducklings ranged from 0.15 (95% CI = 0.04-0.27) to 0.24 (95% CI = 0.13-0.38) and was 0.21 (95% CI = 0.15-0.28) for 1996–1999. Our overall estimate of brood survival was 0.64 (n = 91; 95% CI = 0.54-0.73). At TTRW, survival of 129 radiomarked ducklings was 0.29 in 1998 (95% CI = 0.20-0.41) and 1999 (95% CI = 0.13-0.45) and was 0.29 (95% CI = 0.20-0.40) for 1998–1999. Our overall estimate of brood survival was 0.71 (n = 38; 95% CI = 0.56-0.85). At NNWR, models that included all predictor variables best explained variation in duckling survival. Akaike weight (w_i) for the best model was 0.81, suggesting it was superior to other models (<0.01 < w_i < 0.18). We detected 4 competing models for duckling survival at TTRW. Inter-day distance traveled by ducklings was important as this variable appeared in all 4 models; duckling survival was positively related to this variable. Patterns of habitat-related survival were similar at both study areas. Ducklings in broods that used scrub-shrub habitats disjunct from wetlands containing aggregations of nest boxes had greater survival probabilities than birds remaining in wetlands with such nest structures. Managers may increase local wood duck recruitment by promoting availability of suitable brood habitats (e.g., scrub-shrub wetlands) without aggregations of nest boxes that may attract predators and by dispersing nest boxes amid or adjacent to these habitats. We did not determine an optimal density of nest boxes relative to local or regional population goals, which remains important research and conservation needs.     

A Novel Method of Assessing Dominance Hierarchies Shows Nuance,Linearity and Stability in the Dinosaur Ant Dinoponera quadriceps

Many social species with relatively simple societies have dominance hierarchies of individuals, with dominant individuals achieving fitness and subordinate individuals either queuing to obtain fitness or achieving only indirect fitness by helping relatives. Assessing the dominance hierarchy in a social group is generally based upon observing dyadic interactions as and when they occur spontaneously within the whole‐group setting. However, this method can be very time‐consuming because many dyads interact only very rarely, necessitating either extremely long observation periods or many dyadic relationships being unresolved. Here, we report an alternative method using the queenless dinosaur ant Dinoponera quadriceps, which lives in colonies containing tens of individuals. We removed all individuals from their nest and observed the dominance behaviours expressed in isolated dyadic interactions for every pairwise combination of individuals. Individuals showed a classic dominance behaviour in this setting, and the rapid nature of the assay allowed us to observe every dyadic relationship on a weekly basis over 4 weeks. The dominance hierarchies based on these isolated dyadic interactions correlated well with those produced by the conventional method of colony observations. They showed the hierarchies to be highly linear and stable, and also revealed that dominance relationships may extend further down the hierarchy than previously thought. Although highly manipulative, the isolated dyadic interaction method works well and will likely make more feasible the study of other social species in which pairs of individuals can be isolated together.     

Male Mating Tactics in Captive Rhesus Macaques (Macaca mulatta): The Influence of Dominance, Markets, and Relationship Quality

Male mating success in a multimale–multifemale group can depend on several variables: body condition, dominance, coalitions, “friendship,” or an exchange of services for mating access. Exchange patterns may also be determined by market effects or social relationships. We studied the mating tactics of males in a captive, multimale–multifemale group of rhesus macaques and the resulting patterns of mating and paternity to determine the influence of dominance rank, mating markets, and relationship quality on their mating tactics. Male rank was positively related to the total number of copulations and the number of mating partners, but did not explain male mating distribution completely. Moreover, male fertilization success was not related to male rank. Males did not exchange grooming for mating access on the same day and neither the supply nor the rank (as a proxy for quality) of receptive females affected the amount of male grooming, suggesting that market effects did not explain male mating access. However, there was a positive correlation between long-term grooming patterns of both males and females and mating access, indicating that social relationships were important for male mating access. Paternity data revealed that these social relationships were also important for male reproductive success. We conclude that both male rank and male–female “friendship” determined male mating access in these rhesus macaques, but that “friendship” was more important in determining paternity, emphasizing the importance of intersex social bonds in male mating success in multimale primate societies.     

Familiarity and dominance relations among female sooty mangabeys in the Taï National Park

Dominance relationships of female sooty mangabeys have thus far been studied exclusively in captive groups. In captivity, adult females form a stable linear hierarchy as would be expected in species exhibiting strong contest competition. However, the same individuals do not exhibit other aspects of behavior that would be expected where contest competition occurs. For example, they show no kin‐based alliances leading to hierarchies in which the members of each matriline occupy adjacent ranks. The goal of this study was to provide the first data on dominance relationships of sooty mangabey females in their natural environment in the Taï National Park, Ivory Coast. In our study group, adult females formed a linear dominance hierarchy. Aggression over food increased in food patches, as would be expected for species that experience contest competition. Moreover, females formed highly differentiated social relationships, showing particular affinities with females of adjacent rank. Am. J. Primatol. 56:137–153, 2002. © 2002 Wiley‐Liss, Inc.     

Behavioural and physiological correlates of personality in greylag geese (Anser anser)

Personality means suites of correlated behavioural traits, also referred to as “behavioural syndromes” or “personality dimensions”. Across animal taxa similar combinations of traits seem to prevail, which may have proximate foundation in common neuroendocrine mechanisms. Hitherto, these have been rarely studied in intact social settings. We investigated personalities of greylag goose males from a free-roaming flock that shows complex social relationships. In connection with our longitudinal study on the consistency of behavioural and physiological responses to multiple challenges, we asked whether and how single, personality-related behavioural traits correlate with each other to form personality dimension(s). We tested whether these dimensions were related to physiological characteristics that previously showed limited plasticity (heart rate (HR), baseline and stress-induced excreted immuno-reactive corticosterone (BM), and testosterone metabolites levels) and, furthermore, to age, body measures, and dominance rank. Principal-components analysis based on behavioural variables revealed two factors: 51.1% of variability was explained by “aggressiveness” and a further 19.1% by “sociability”. “Aggressiveness” comprised correlated measures of aggression, subordinance, boldness, vigilance, and proximity to the mate. This “aggressiveness” positively correlated with stress-induced BM levels, the HR increase during aggressive interactions, and with dominance rank, which may suggest proximate and functional contingencies of this personality dimension.     

Spatial structure and parental aggression in eider broods

The spatial position of young animals within a brood affects their survival, so that marginal individuals are at greater risk of predation. Spatial brood structuring may be caused by differences in offspring size, age, hunger, or active parental manipulation through aggression. Nepotistic manipulation of brood structure would confer fitness benefits for parents accepting nondescendant young. However, insufficient kin recognition has often been considered to preclude such nepotism in birds, particularly in precocial waterfowl. We explored the spatial structure of ducklings within broods of eiders, Somateria mollissima, a seaduck with frequent brood amalgamation. We compared the distribution of ducklings of different origin relative to reference females whose kinship to the ducklings was known. We also observed female aggression towards ducklings, to evaluate the role of parental manipulation of brood structure. We found a nonrandom distribution of ducklings within broods; a female's own young were on average closer to her than unrelated young were. We also found evidence for parental nepotism: whether the brood contained unrelated young was the strongest predictor of female aggression towards ducklings. The spatial position, hatch weight and relative size of ducklings showed no significant correlations with each other, suggesting that active parental manipulation may be needed to explain the observed spatial structure. Our study conflicts with previous anecdotal evidence suggesting that brood amalgamation in eiders results in the disintegration of parent-offspring bonds, preventing parental exploitation of nondescendant young. It also opens up the possibility that the spatial position of ducklings depends on their mother's status in the female dominance hierarchy. Copyright 2003 The Association for the Study of Animal Behaviour. Published by Elsevier Ltd. All rights reserved.      

Female philopatry and dominance patterns in wild geladas

Cercopithecines have a highly conserved social structure with strong female bonds and stable, maternally inherited linear dominance hierarchies. This system has been ascribed to the pervasiveness of female philopatry within the typical multi-male, multi-female social groups. We examined the relationship between female philopatry, dominance hierarchies, and reproduction in geladas (Theropithecus gelada), a species with an unusual multi-leveled society. During a 4-year field study on a wild population in the Simien Mountains National Park, Ethiopia, we observed 14 units across two bands of geladas that underwent a number of events, such as male takeovers and female deaths, which could potentially disrupt female relationships and unit structure. First, we corroborate earlier reports that gelada females are natally philopatric: we observed no interunit migrations, and the female mortality rate was comparable to that of philopatric baboons (suggesting all female disappearances were indeed deaths). Second, contrary to previous reports, data from this long-term study show that geladas exhibit the linear and stable dominance hierarchies typical of other Cercopithecines. Moreover, female ranks appear to be maternally inherited. Third, we found no evidence that alpha females aggressively target the lowest ranking individuals, nor did rank confer clear reproductive advantages to dominant females within our 4-year observation period. As such, geladas fit the allostatic load model [Goymann & Wingfield, Animal Behaviour 67:591-602, 2004]. Our study confirms the importance of female philopatry in the kin-based Cercopithecine dominance system.     

Seasonal rank changes for adolescent and subadult natal males in a free-ranging group of rhesus monkeys

Changes in dominance rank for adolescent and subadult natal males in a semi-free-ranging rhesus macaque group were seasonal. Three 4-year-old natal males of the highest-ranking matriline occupied high ranks in the adult male dominance hierarchy during the premating period. In the mating season they dropped in rank, and this decrement was related to a concomitant drop in their alliances. After the mating season, these males rose in rank along with their two 3-year-old kin to occupy ranks 2–5 and 7–8 in the adult male dominance hierarchy. Three-year-old natal males of matrilines lower ranking than first did not become integrated into the hierarchy at this time.     

Food Competition and Linear Dominance Hierarchy Among Female Chimpanzees of the Taï National Park

Dominance rank in female chimpanzees correlates positively with reproductive success. Although a high rank obviously has an advantage for females, clear (linear) hierarchies in female chimpanzees have not been detected. Following the predictions of the socio-ecological model, the type of food competition should affect the dominance relationships among females. We investigated food competition and relationships among 11 adult female chimpanzees in the Taï National Park, Côte d'Ivoire (West Africa). We detected a formal linear dominance hierarchy among the females based on greeting behaviour directed from the subordinate to the dominant female. Females faced contest competition over food, and it increased when either the food was monopolizable or the number of competitors increased. Winning contests over food, but not age, was related to the dominance rank. Affiliative relationships among the females did not help to explain the absence of greetings in some dyads. However comparison post hoc among chimpanzee study sites made differences in the dominance relationships apparent. We discuss them based on social relationships among females, contest competition and predation. The cross-site comparison indicates that the differences in female dominance hierarchies among the chimpanzee study sites are affected by food competition, predation risk and observation time.     

Social Dominance Hierarchies and Phenotypic Correlates of Dominance in Captive Groups of the Grey‐cheeked Fulvetta Alcippe morrisonia in Taiwan

Dominance hierarchies usually form quickly among avian foraging groups because they are beneficial to most individuals by reducing conflict. Several characteristics that correlate with dominance rank have been identified in birds, but most of these conclusions rely on studies of temperate species. Hence, we studied whether captive group members of a subtropical species, grey‐cheeked fulvetta Alcippe morrisonia, form social dominance hierarchies when competing for food during the non‐breeding season. We also investigated whether sex, age, body condition and fat score were related to an individual's dominance rank which was established by counting aggressive interactions in six captive groups of nine individuals each. In all groups, linear dominance hierarchies were formed whereby yearlings dominated over adult birds, and individuals with a better body condition were also more dominant, while sex and fat score had no discernable effect. Male yearlings had significantly higher body masses and body condition indices than male adults, while female yearlings had significantly higher body masses, body condition indices and fat scores than female adults. However, there were no significant differences between male and female yearlings or adults for any of these variables. We suggest possible reasons for the dominance of yearlings, such as captive conditions or the higher body weight of yearlings.     

Survival of common goldeneye Bucephala clangula ducklings in relation to weather, timing of breeding, brood size, and female condition

The survival of common goldeneye Bucephala clangula ducklings during their first week of life was studied in relation to hatching date, brood size, female condition, and weather (temperature and precipitation) during the first week post-hatch by using data from radio-marked females and their broods. Also, age-specific variation in the survival of the young was determined until fledging (over 50 d of age). Survival was lowest in the first week after hatching. Hatching date, brood size, and first week temperature and precipitation were poor predictors of duckling survival during the first week after nest exodus. Instead, the ducklings of females in a better body condition survived better in their first week of life. The results suggest that weather does not have a direct effect on downy ducklings' survival, but the condition of the female seems to be an important determinant of the survival of common goldeneye ducklings.     

The benefit of large broods in barnacle geese: a study using natural and experimental manipulations

1. In precocial birds, where the young feed themselves, the costs and benefits of brood size are still poorly understood. An experimental manipulation of brood size was employed to examine the effects of brood size on both parents and young in a wild population of barnacle geese [ Branta leucopsis (Bechstein)] during brood-rearing on Svalbard.
2. Social dominance of the family unit, the amount of vigilance behaviour of the parents, the growth of the goslings in the family unit and an index of body condition for female parents during moult were all positively correlated with brood size.
3. When brood size changed as a result of natural events (i.e. predation or adoption) or experimental manipulation, rates of dominance, parental vigilance, gosling growth and female parent condition changed in a similar direction to the observed relation between the variable and brood size in unchanged broods.
4. After fledging, the fast-growing goslings in large broods survived better during autumn migration, while there was no apparent net cost in survival or next-year breeding for the parents.
5. Via a direct effect of brood size on dominance of the family unit, large broods were beneficial for both parent and young in a situation where there was strong intraspecific competition for the available food resources.
6. This study provides a clear demonstration of a causal relationship between brood size and various components of both gosling and adult fitness and is of direct relevance to the phenomenon of adoption and the evolution of brood size in this species.     

Longitudinal changes of dominance rank among the females of the Koshima group of Japanese monkeys

The changes of dominance rank among female Japanese monkeys of the Koshima group over a period of 29 years from 1957 were studied. The dominance rank order was relatively stable in the early population growing phase, while large scale-changes of dominance rank order occurred successively in the phase of population decrease brought about by the severe control of artificial feeding after 1972. Nevertheless, the rank order of several females of the highest status was stable. Furthermore, the reproductive success of these highest status females was high (Mori, 1979a;Watanabe et al., in prep.). Divergence of the dominance rank order fromKawamura's rules (Kawamura, 1958) was observed in the following respects: (1) Some females significantly elevated their rank depending on the leader males. (2) If mothers died when their daughters were still juveniles or nulliparous, the dominance rank of some of these offspring females was significantly lower than the mother's one. However 55% of daughters which lost their mothers at a young age inherited the mother's rank. (3) Dominance among sisters whose mother had died when at least one of the daughters was under 6 years old followed the rule of youngest ascendancy in 60% (Kawamura, 1958), and in 80% when both of the daughters were nulliparous at the mother's death. The mean rate of aggressive interactions for each female with subordinates to her was calculated by dividing the total aggressive interactions between the female in question and her subordinates by the number of subordinate females to the female in question. A female which showed a high rate of aggressive interactions with her subordinates was categorized as an “Attacker”, and a female showing a lower rate was categorized as a “Non-attacker”. Similarly, categories of “Attacked”, and “Non-attacked” were distinguished by using the rate of aggressive interactions with dominant females. Several females which were once categorized in one category in a year were repeatedly categorized in the same category over different years. The “Attacked” tended to be females of higher rank, and “Non-attackers” tended to be females of lower rank. “The second-higher-status females”, were “Attacked”, and their rank was unstable. In particular, females of lower rank within the lineage of the highest rank suffered this kind of severe status. Most of the daughters of these females showed a sharp drop of rank, and died when they were still at a young age, i.e. “the second-higher-status females” displayed low fitness. “Non-attackers” were significantly “Non-attacked”; i.e. they were females which showed a non-social attitude. Females which underwent a drop of rank tended to be “Non-attackers”. The most important factor which determined the females' rank was the memory of their dominance relations under the influence of their mother [dependent rank (Kawai, 1958)] in their early life during development. This finding corresponds well with the results in baboons obtained byWalter (1980); the target females of aggressive interactions by adolescent females were determined by the rank of the mothers when these adolescent females were born.     

Dominance and its Behavioral Measures in a Captive Group of Bonobos (Pan paniscus)

We investigated the existence of a social dominance hierarchy in the captive group of six adult bonobos at the Planckendael Zoo. We quantified the pattern of dyadic exchange of a number of behaviors to examine to what extent each behavior fits a linear rank order model. Following de Waal (1989), we distinguish three types of dominance: agonistic dominance, competitive ability and formal dominance. Fleeing upon aggression is a good measure of agonistic dominance. The agonistic dominance hierarchy in the study group shows significant and strong linearity. The rank order was: 1. female (22 yr), 2. female (15 yr)., 3. male (23 yr.), 4. female (15 yr.), 5. male (9 yr.), 6. male (10 yr.). As in the wild, the females occupy high ranks. There is prominent but nonexclusive female agonistic dominance. Teeth-baring does not fulfil the criteria of a formal submission signal. Peering is a request for tolerance of proximity. Since its direction within dyads is consistent with that of fleeing interactions, it is a useful additional measure to determine agonistic ranks in bonobos. In competitive situations, the females acquire more food than other group members do. The rank obtained from access to food resources differs from the agonistic rank due to female intrasexual social tolerance, expressed in food sharing. We typify the dominance styles in the group as female intrasexual tolerance and male challenging of rank differences. The agonistic rank order correlates significantly with age and has a strong predictive value for other social behaviors.     

Male dominance and genetically determined reproductive success in the mandrill (Mandrillus sphinx)

Darwin referred to the adult male mandrill (Mandrillus sphinx) as the most brightly coloured of all mammals, citing the brilliant red and blue pigmentation of the face, rump, and genitalia as extreme examples of evolution by sexual selection. Considerable controversy exists concerning possible effects of sexually selected phenotypes via intermale competition on reproductive success. Behavioural and genetic studies of a large, semi-free ranging mandrill colony in Gabon have now demonstrated that clear-cut relationships exist between male secondary sexual development, social dominance, copulatory behaviour, and reproductive success in this primate species. Two morphological variants of adult male were identified; “fatted” males, with maximum secondary sexual coloration, which occupied dominant positions in the social group, and “non-fatted” males, with muted secondary sexual adornments, smaller testes and lower plasma testosterone levels, which lived as peripheral/solitary individuals. DNA fingerprinting analyses on infants born over five successive years showed that only the two most dominant, fatted males in the group had fathered offspring. Throughout the annual mating season these males attempted to mate-guard and copulate with females during periods of maximal sexual skin tumescence. Male rank and mating success were strongly positively related and the alpha male sired 80 – 100% of the resulting offspring during three consecutive years. Non-fatted adult males and group associated subadult males engaged in infrequent, opportunistic matings and did not guard females. Loss of alpha status resulted in a fall in reproductive success, but the effect was gradual; the deposed alpha male continued to father 67% and 25% of infants born during the next two years. Thus, whilst claims that male dominance determines mating success and paternity in primates have caused considerable debate, these results on mandrills provide unequivocal evidence for the existence of such effects.