A Young Manakin Knows His Place: Evidence for an Age‐Graded Dominance Hierarchy Among Long‐Tailed Manakins

In lek‐breeding systems where many males gather at display sites, males benefit from the establishment of dominance hierarchies to reduce intrasexual aggression and the associated risk of injuries. Long‐tailed manakins (Chiroxiphia linearis) exhibit an exploded lek‐breeding system wherein the two top‐ranking males at each display site team up to perform elaborate coordinated courtship displays for females. Young males undergo delayed plumage maturation whereby they acquire distinct pre‐definitive plumage patterns each year until they attain definitive plumage in their fifth year. This unique characteristic is thought to have evolved as a status‐signalling mechanism to aid in the establishment of an age‐graded dominance hierarchy in which older males are dominant to younger males. Previous research has shown evidence for such a dominance hierarchy among alpha and beta males; however, the presence of this hierarchy among males of other age classes has never been quantified. In this study, we investigated the presence of an age‐graded dominance hierarchy by determining whether older males direct more aggressive behaviours towards younger males. We also investigated whether status signalling is less clear within age classes than between age classes, by determining whether males within the same age class exhibit more aggression towards each other. We found that older males performed aggressive behaviours towards younger males much more frequently than younger males performed aggressive behaviours towards older males. We also found that some aggressive interactions occurred between males within the same age class more frequently than between males from different age classes. Our study provides some evidence for an age‐graded dominance hierarchy among male long‐tailed manakins of all age classes and also provides some support for the status‐signalling hypothesis. However, further research is needed to conclusively establish the presence of a linear dominance hierarchy among younger male manakins. This research may help us better understand the evolution of complex hierarchical systems in animals.     

Social organization of the spotted hyaena Crocuta crocuta. II. Dominance and reproduction

A 4-year study of the social organization of spotted hyaenas in a clan of 60–80 individuals showed that there is a separate dominance hierarchy within each sex. One female and her descendants dominated all others; matrilineal rankings were stable over time because maternal rank is inherited. Cubs of higher ranking females were able to feed at kills in competition with adults more successfully than other cubs, and male offspring of the alpha female were the only males able to dominate adult females. The mating system is highly polygynous: only the behaviourally dominant male was seen to mate, though all other resident males regularly courted females. Among females, there was no correlation between reproductive success and age, size, or social rank. It is postulated that the unusually aggressive sons of the alpha female would probably be highly successful competitors in the context of a polygynous mating system. A primary consequence of female dominance over males is that females and their young have priority of access to food in a highly competitive feeding situation. This competition may have been the selective force that produced female dominance and the associated syndrome of female virilization that is characteristic of the species. Cooperation among related females may be the basis for the matrilineal system, as has been suggested for certain primate species.     

Life history in male mandrills (Mandrillus sphinx): physical development, dominance rank, and group association

We assess life history from birth to death in male mandrills (Mandrillus sphinx) living in a semifree-ranging colony in Gabon, using data collected for 82 males that attained at least the age of puberty, including 33 that reached adulthood and 25 that died, yielding data for their entire lifespan. We describe patterns of mortality and injuries, dominance rank, group association, growth and stature, and secondary sexual character expression across the male lifespan. We examine relationships among these variables and investigate potential influences on male life history, including differences in the social environment (maternal rank and group demography) and early development, with the aim of identifying characteristics of successful males. Sons of higher-ranking females were more likely to survive to adulthood than sons of low-ranking females. Adolescent males varied consistently in the rate at which they developed, and this variation was related to a male's own dominance rank. Males with fewer peers and sons of higher-ranking and heavier mothers also matured faster. However, maternal variables were not significantly related to dominance rank during adolescence, the age at which males attained adult dominance rank, or whether a male became alpha male. Among adult males, behavior and morphological development were related to a male's own dominance rank, and sons of high-ranking females were larger than sons of low-ranking females. Alpha males were always the most social, and the most brightly colored males, but were not necessarily the largest males present. Finally, alpha male tenure was related to group demography, with larger numbers of rival adult males and maturing adolescent males reducing the time a male spent as alpha male. Tenure did not appear to be related to characteristics of the alpha male himself.     

Perseverance and food sharing among closely affiliated female chimpanzees

Chimpanzees (Pan troglodytes) have been frequently observed to share food with one another, with numerous hypotheses proposed to explain why. These often focus on reciprocity exchanges for social benefits (e.g., food for grooming, food for sex, affiliation, kinship, and dominance rank) as well as sharing based on begging and deterring harassment. Although previous studies have shown that each of these hypotheses has a viable basis, they have only examined situations in which males have preferential access to food whereby females are required to obtain the food from males. For example, studies on male chimpanzee food sharing take advantage of successful crop-raids and/or acquisitions of meat from hunting, situations that only leave females access to food controlled by male food possessors. This begs the question how and with whom might a female chimpanzee in sole possession of a high-quality food item choose to share? In two large captive groups of chimpanzees, we examined each of the hypotheses with female food possessors of a high-quality food item and compared these data to a previous study examining food transfers from male chimpanzees. Our results show that alpha females shared significantly more with closely affiliated females displaying perseverance, while kinship and dominance rank had no effect. This positive interaction between long-term affiliation and perseverance shows that individuals with whom the female possessor was significantly affiliated received more food while persevering more than those with neutral or avoidant relationships towards her. Furthermore, females with avoidant relationships persevered far less than others, suggesting that this strategy is not equally available to all individuals. In comparison to the mixed-sex trials, females chose to co-feed with other females more than was observed when the alpha male was sharing food. This research indicates that male and female chimpanzees (as possessors of a desired food item) share food in ways influenced by different factors and strategies.     

Sexual behavior of adult male chimpanzees of the Mahale Mountains national park,Tanzania

This study, based on 687 hr of focal observations, aims to describe overall patterns of the sexual behavior of the adult male chimpanzees of the Mahale Mountains, to compare the results with previous reports, and to explain the variations between studies. Genital inspection of cycling females by adult males was eight times as frequent as that of lactating females, and twice as frequent as that of pregnant females. Inspection of the genitals of cycling females increased dramatically 7–10 days before the onset of maximal swelling and gradually decreased as the day of ovulation approached. Adult males likely obtained information on the attractivity of females by inspecting their genitals. Mating was usually initiated by male courtship and followed by pelvic thrusts in a dorsoventral posture, performed on, rather than above, the ground, which continued for 7 s. on average, and was typically followed by female squeaking and darting from the male, or by the male grooming the female. Higher-ranking males mated with females in the peri-ovulatory period more frequently than did lower-ranking males. In particular, two alpha males mated with such females more often than did any other adult males. A male who interfered with a mating pair was dominant over the mating male in other agonistic contexts. The duration of intromission was correlated with neither dominance rank nor age. However, when an adult male declined in rank from alpha in 1991 to third in 1992, he showed a significantly shorter duration of intromission. This indicates that for a particular male, the alpha rank guaranteed longer duration of intromission. Allies of alpha males tended to mate with peri-ovulatory females more frequently than expected from their low dominance ranks. The number of mating partners was not correlated with male dominance rank, but was sometimes negatively correlated with male age. Females were significantly more likely to emit a copulatory squeak when mating with younger, rather than older, adult males. Male dominance rank and the rate of female copulatory squeaking were not correlated. Weaning infants regularly interfered with their mothers' mating. Occasionally, unrelated adolescent males and rarely females pushed themselves in between copulating adults. Female choice was indicated when they performed a “penis erection check” or took the initiative in courtship, or on the other hand showed strong reluctance to mate with particular males. Young adult males more often received erection checks than did prime males, while none of the three old adult males did. Courtship initiated by estrous females was not directed to two of the oldest males, the exception of which was the alpha male. The oldest males, except for the alpha, were consistently avoided by many estrous females, both young and old. In response to female reluctance, males behaved violently, however, this was not effective, because other more dominant males came to rescue the female. Neither courtship nor mating was seen between mature sons and their mothers, nor between brothers and sisters.     

Effects of Removal of Successful Males in a Fallow Deer Lek

Removal of 5 of the 8 most successful males in a fallow deer lek between breeding seasons led to an increase in fighting rate of males and a decrease in mating rate of both sexes. The increase in fighting was likely due to disruption of the social hierarchy. The lower mating rate of females was not due to disruption of copulations by territorial males, nor did it appear to result from disturbance from fights. We suggest that difficulties in mate choice related to an unstable male hierarchy resulted in females either increasing their length of stay in the lek, visiting the lek more than once before mating, or leaving the lek without mating.     

Biochemical and hormonal correlates of dominance and social behavior in all-male groups of squirrel monkeys (Saimiri sciureus)

This study examined the relationship between dominance rank and several physiological and behavioral measures in stable, captive, all-male squirrel monkey groups. Four groups, each containing three males, were observed for 12 weeks during the breeding season. Relative dominance ranking among males in each group was based on the direction of agonistic behaviors displayed. For each subject, whole blood serotonin (WBS), plasma testosterone (T), and cortisol (C) were sampled 4 to 6 times over the course of the study. Samples were separated by 1- to 2-week intervals. Each group had a stable linear dominance hierarchy. WBS had small intraanimal variance and was positively related to dominance rank. In contrast, T and C were highly variable within subjects and were unrelated to dominance rank. Among alpha males, concentrations of T and C were positively correlated, and WBS and C were negatively correlated. The effect of dominance rank on the relationship between within-animal fluctuations in WBS, T, and C and behavior was also assessed. In dominant, but not subordinate subjects, intraanimal fluctuations in WBS correlated with agonistic behavior initiated, and fluctuations in C and T correlated with huddling. In beta and gamma males, C was related to agonism received, and in gamma males to food stealing received. Dominance status also affected endocrine response to the stress of capture but not the rate of sneezing. Sneezing was positively correlated with T concentrations irrespective of dominance rank. These results extend the association between WBS and dominance rank previously reported in Old World monkeys to a New World monkey species, support previous suggestions that mean T and C titers are not reflective of dominance rank in well-established groups, and indicate that dominance rank affects adrenocortical response to the stress associated with capture and anesthesia.     

The changes in the dominance hierarchy over time of a complete field-captured colony of Cryptomys hottentotus hottentotus

The common mole-rat, Cryptomys h. hottentotus , is a social subterranean rodent occurring in colonies in which one female and one to three males are involved in reproduction and the remaining colony members are non-reproductive. Within each sex the reproductive animals are usually the largest and most dominant animals.
The dominance hierarchy amongst a field-captured colony was linear ( h = 0.95, calculated from Landau's linearity index) soon after capture. The non-reproductive females were ranked low in the dominance hierarchy; many were subordinate to non-reproductive males. The order of capture of mole-rats was not related to the position in the dominance hierarchy. The hierarchy became non-linear ( h = 0.56) after six months in captivity during which two juvenile animals became adult. The breakdown in the hierarchy may result from the lack of opportunity in captivity for animals to disperse and establish satellite colonies, or from colony members becoming co-dominant in the hierarchy as a result of a rise in rank by young animals.
Dominant mole-rats are involved in a greater proportion of interactive behaviours than subordinates. Popularity studies show that females tend to be more popular animals than males. The largest reproductive male was the least popular animal in the first study, whereas a beta male was the least popular animal in the second study period. The reproductive female was the most popular in both periods.     

Social correlates of variation in urinary cortisol in wild male bonobos (Pan paniscus)

Cortisol excretion in males of group living species is often associated with social rank and competition for oestrous females. Rank-related patterns of cortisol levels can be used to study mechanisms of rank maintenance and costs associated with mate competition. Bonobos (Pan paniscus) are interesting because males form a linear dominance hierarchy but are not dominant over females and therefore aggressive male-male competition over access to females alone is not considered to be a successful reproductive strategy. In this study on social correlates of urinary cortisol in wild male bonobos, we investigated the relationship between cortisol levels and several aspects of mate competition, including male rank, aggression rates, and association time with oestrous females. We found that cortisol levels correlated positively with dominance rank when oestrous females were present, but not when they were absent. This result is consistent with the idea that aggressive behaviour plays a minor role in maintenance of high rank. While aggression received from males and females explained within-individual variation in cortisol levels, it was the time spent in association with oestrous females that best explained between-individual variation in male cortisol levels. The observed increase in male cortisol may be associated with spatial proximity to oestrous females and could result from anticipated aggression from other group members, reduced feeding time in the males, or a combination of both.     

The Social Implications of traditional use of lek sites in the ruff Philomachus pugnax

In almost every letting vertebrate, adult males use traditionallek sites. This paper attempts to explain this phenomenon byinvestigating the social implications of traditional use oflek sites. A population of ruffs Philomachus pugnax was observedduring six breeding seasons, and lek stability, site fidelity,male copulatory success, and male dominance relationships werestudied. Ninety percent of the territorial males were site-faithfulwithin and between years, and male dominance, copulatory success,and order of territory establishment were all inter-correlated.Males that were relatively dominant and successful as first-yearterritorials were more likely to return to the area, establishedthemselves even earlier in their second year, and rose in successrank. The frequency of disrupted copulations was very low, despitethe tightly clustered territories on leks. This suggests thatwell-developed dominance relationships exist between males.Site-faithful males are more likely to acquire accurate informationabout the competitive abilities of other males, and such informationis probably necessary for stable dominance relationships todevelop. Such relationships reduce the intensity of male aggression,thereby reducing the risk of male injury. High levels of maleaggression also deter females. Furthermore, stable dominancerelationships enables low-and medium-ranked males to remainon the leks while waiting for the top males to drop out. Thehigh site-fidelity of lekking males, thus, is suggested to haveevolved to facilitate the establishment of stable dominancerelationships, which are beneficial to all territorial males,irrespective of rank     

Stress, social behaviour, and secondary sexual traits in a male primate

We examined variation in glucocorticoid levels in the mandrill, a brightly coloured primate species, to identify major social influences on stress hormones, and investigate relationships among glucocorticoid levels, testosterone and secondary sexual ornamentation. We collected a total of 317 fecal samples for 16 adult male mandrills over 13 months, including mating and non-mating periods and periods of both dominance rank stability and instability, and compared fecal glucocorticoid levels with dominance rank, rank stability, presence of receptive females, gastro-intestinal parasite infection, fecal testosterone and facial red coloration. Glucocorticoid levels did not vary systematically with dominance rank, but increased when the dominance hierarchy was unstable, and increased in the presence of receptive females. The relationship between dominance rank and glucocorticoid levels changed direction according to the stability of the dominance hierarchy: glucocorticoid levels were higher in subordinate males under stable conditions, but under conditions of instability higher ranking males had higher glucocorticoid levels. The influence of dominance rank also interacted with the presence of receptive females: glucocorticoids were higher in dominant males than in subordinates, but only during mating periods, suggesting that dominant males are more stressed than subordinates during such periods. These findings support previous studies showing that the relationship between glucocorticoids and dominance rank in male baboons is dependent on the social environment. We also found that males with higher glucocorticoids suffered a higher diversity of gastrointestinal parasite infection, in line with evidence that glucocorticoids suppress the immune system in other species. However, we found no support for the stress-mediated immunocompetence handicap hypothesis for the evolution of condition-dependent ornaments: glucocorticoid and testosterone levels were positively related, rather than the negative relationship predicted by the hypothesis, and we found no relationship between red colour and glucocorticoid levels, suggesting that glucocorticoids do not play a role in translating social conditions or physical health into ornament expression in this species.     

Male Social Behavior and Dominance Hierarchy in the Sulawesi Crested Black Macaque (Macaca nigra)

In a 6-week study of the social behavior of wild Sulawesi crested black macaques (Macaca nigra), we found a linear and transitive dominance hierarchy among the six adult males in one social group. Dominance rank, as determined by the direction of supplantations, correlated strongly with percentage of time near more than four neighbors, frequency of grooming received from adult females, and percentage of time with an adult female as nearest neighbor. These results suggest that high-ranking males are socially attractive. Adult females sexually solicited high-ranking males more often than low-ranking males, but frequency of copulation was not correlated with dominance rank. Frequency and intensity of aggression between males are strongly correlated with rank distance, but aggression toward females was greatest for mid-ranking males. Males of all rank displayed significantly more aggression toward sexually receptive females than toward females in other estrous states. These data indicate that male Sulawesi crested black macaques display a social organization similar to that reported for multimale groups in other macaque species rather than the egalitarian social organization described for female Sulawesi macaques.     

Queuing and queue-jumping: long-term patterns of reproductive skew in male savannah baboons, Papio cynocephalus

In many animals, variance in male mating success is strongly correlated with male dominance rank or some other measure of fighting ability. Studies in primates, however, have varied greatly in whether they detect a relationship between male dominance rank and mating success. This variability has led to debate about the nature of the relation between rank and mating success in male primates. We contribute to the resolution of this debate by presenting an analysis of the relationship between dominance rank and male mating success over 32 group-years in a population of wild savannah baboons. When data were pooled over the entire period, higher-ranking males had greater access to fertile females. However, when we examined successive 6-month blocks, we found variance in the extent to which rank predicted mating success. In some periods, the dominance hierarchy functioned as a queue in which males waited for mating opportunities, so that rank predicted mating success. In other periods, the queuing system broke down, and rank failed to predict mating success when many adult males were in the group, when males in the group differed greatly in age, and when the highest-ranking male maintained his rank for only short periods. The variance within this single population is similar to the variance observed between populations of baboons and between species of primates. Our long-term results provide strong support for the proposition that this variance is not an artefact of methodological differences between short-term studies, but is due to true variance in the extent to which high-ranking males are able to monopolize access to females. Copyright 2003 Published by Elsevier Science Ltd on behalf of The Association for the Study of Animal Behaviour.     

The effects of temporary removal of the alpha male on the behavior of subordinate male vervet monkeys

The effects of temporary removal of the alpha male on the behavior of subordinate adult male vervet monkeys were evaluated. Twelve subordinate males from six captive multimale, multifemale vervet monkey groups were observed in two conditions: when all group members were present and when the alpha male was temporarily removed from each group. In the absence of the alpha male, subordinate males initiated more affiliative behavior and increased the amount of time spent in proximity to females but their rates of aggression were unaltered. Increased affiliative behavior was selectively directed to high-ranking females and their offspring. Under removal conditions, subordinate male behavior did not resemble that of alpha males in intact conditions: they differed in their proximity to and affiliative behavior towards other group members. In the absence of the alpha male, females increased their aggression towards subordinate males. These observations suggest that the presence of alpha males strongly inhibits subordinate males' behavior. When the constraints of the alpha male's presence are removed, subordinate males rapidly engage in behavior that may enhance their likelihood of attaining high rank. In combination with prior studies, the data also indicate that the behaviors involved in the maintenance of high rank by alpha males differ from those subordinates use to acquire dominance. Finally the current study supports the view that aggression by female vervets may be highly influential in determining male ascendency to dominant rank.     

Benefits and Costs of Dominance in the Angelfish Centropyge bicolor

Social groups are often structured by dominance hierarchies in which subordinates consistently defer to dominants. High‐ranking individuals benefit by gaining inequitable access to resources, and often achieve higher reproductive success; but may also suffer costs associated with maintaining dominance. We used a large‐scale field study to investigate the benefits and costs of dominance in the angelfish Centropyge bicolor, a sequential hermaphrodite. Each haremic group contains a single linear body size‐based hierarchy with the male being most dominant, followed by several females in descending size order. Compared to their subordinate females, dominant males clearly benefited from disproportionately high spawning frequencies, but bore costs in lower foraging rates and greater aggressive defence of their large territories. Within the female hierarchy, more dominant individuals benefited from higher spawning frequencies and larger home ranges, but displayed neither higher foraging rates nor spawn order priority. However, dominance in females was also linked to aggressiveness, particularly towards immediate subordinates, suggesting that females were using energetically costly aggression to maintain their high rank. We further showed by experimentally removing dominant females that the linear hierarchy was also a social queue, with subordinates growing to inherit higher rank with its attendant benefits and costs when dominants disappeared. We suggest that in C. bicolor, the primary benefit of high rank is increased reproductive success in terms of current spawning frequency and the prospect of inheriting the male position in the near future, which may be traded off against the cost of aggressively defending rank and territory.     

Establishment of a free-ranging colony of stumptail macaques (Macaca arctoides): Social relations I

Data on the social behaviour of a free-ranging troop of Stumptail macaques are reported for a period starting of August 15th to November 7th of 1974. Positive interactions take a greater part in the daily life of the troop than negative interactions. Individuals order themselves in an essentially linear dominance hierarchy and offspring rank immediately below their mother. Agonism and displacement are good predictors of dominance rank. Dominant individuals are a strong source of attraction for subordinates and the alpha male is a powerful social magnet for immatures. Grooming was found to be correlated with dominance rank and individuals holding adjacent ranks tend to interact positively with one another. To gain access to dominant females, subordinates direct a great deal of positive behaviour to the female’s youngest offspring. As social distance decreases the selection of a social partner becomes more strict and tends toward adjacency. However, factors such as motherhood, genealogical ties, and special relationships with non adjacent individuals, can override the tendency. The direction and quantity of male-care displayed by the adult male and by juvenile males toward younger individuals was found to be influenced by the troop’s dominance patterns. Sex differences were found to exist in male-care received and a negative association was found with the age of the receiver. Adult females are the main regulators of social relations and their rank and friendships provide their offspring with the social format, before puberty, in which they develop socially. The alpha male is the group protector, the performer of policing, an important element in the immatures’ social development, and a powerful source of attraction for all troop members.     

Changes in the dominance rank and reproductive behaviour of male bonnet macaques (Macaca radiata)

The dominance rank of male bonnet macaques influenced their associations and sexual interactions with cycling and conceiving females. Only high-ranking males formed exclusive associations with high-ranking females, and high-ranking males copulated with these females more often than did other males. Changes in male dominance rank were directly related to changes in male mating behaviour, as males who rose in rank spent increasing amounts of time in exclusive proximity to conceiving high-ranking females, and males whose ranks declined spent decreasing amounts of time in proximity to such females. The year of the study in which dominance relationships were most stable was also the year in which the top-ranking male most successfully monopolized access to conceiving high-ranking females.     

Female mate choice across spatial scales: influence of lek and male attributes on mating success of blue-crowned manakins

Lekking males compete for females within and among leks, yet female choice is expected to work differently at each of these spatial scales. We used paternity analyses to examine how lek versus male attributes influence mate choice in the blue-crowned manakin Lepidothrix coronata. We tested the hypotheses that females prefer (i) to mate at larger leks where a larger number of potential mates can be assessed, (ii) to mate with unrelated or highly heterozygous males expected to produce high-quality offspring, (iii) to mate with males that display at higher rates, and that (iv) display honestly reflects male genetic quality. Our results show that (i) males at larger leks are not more likely to sire young, although females nesting close to small leks travel further to reach larger leks, (ii) siring males are not less related to females or more heterozygous than expected, (iii) within a lek, high-display males are more likely to sire young, and (iv) both male heterozygosity and display rate increased with lek size, and as a result display does not reliably reflect male genetic quality across leks. We suggest that female mate choice in this species is probably driven by a Fisherian process rather than adaptive genetic benefits.     

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.     

Mating preference of female zebrafish, Danio rerio, in relation to male dominance

Mating success tends to be skewed toward dominant males, thoughfemale mate preferences may not always correlate with male dominance.In this study, we investigated the mating preferences of femalezebrafish, Danio rerio, in the absence of male–male competition.We paired females sequentially with males of known dominancerank, using a nested, repeated measures design, with egg productionas a measure of female mate preference. We predicted that femaleswould spawn more frequently and produce larger clutches whenpaired with males of higher dominance rank. We found significantdifferences among females in the size of clutches produced andamong males in the size of clutches received, but these differenceswere independent of male dominance rank. Male body size wasnot related to either dominance rank or clutch size received.These results indicate that females vary clutch size in relationto the males with which they are paired but that they do notfavor dominant males. Thus, male competition may normally overridefemale mate preference in zebrafish.