Rank-dependent hoarding effort in willow tits (<Emphasis Type="Italic">Parus montanus</Emphasis>): a test of theoretical predictions

Many birds and mammals store energy as hoarded food supplies. A supply of stored food is beneficial during periods when food is scarce, but building up and managing such a supply also entails costs. The optimal number of caches will be reached when the net benefit is at its maximum. If dominants can steal more stored food from subordinates than the other way around, the optimum will differ between these categories. A previous theoretical model of hoarding in groups with dominant and subordinate members produced three testable predictions: (1) hoarders should store more food as anticipated future conditions get worse; (2) subordinate flock members should store more food than dominants; and (3) dominants should increase hoarding relatively more than subordinates as conditions get worse. Here we present a field experiment on willow tits (Parus montanus) designed to test these predictions. We found support for all three. Hoarding increased as conditions got worse, subordinates stored at a higher rate than dominants, and dominants increased their hoarding effort relatively more than subordinates as conditions worsened. These results support the incorporation of information on dominance and food availability into models predicting food storage behaviour.Communciated by J. Dickinson     

The costs and benefits of territory sharing for the Caribbean coral reef fish,Scarus iserti

Summary Along the Caribbean coast of Panama, groups of unrelated female striped parrotfish, Scarus iserti, co-defend a common feeding territory. Field manipulations of group size and composition were performed to examine the benefits and costs accrued by dominant fish within these shared territories. Dominant fish benefit from the presence of relatively large subordinates because they share in the defense of the territory. Removals of these fish caused increases in defense time and decreases in feeding time for dominant group members. Dominants benefit from the presence of small subordinates because they increase the foraging efficiency of dominants. Removals of smaller subordinates caused reductions in the feeding time of dominant fish, although no changes in defense time occurred. Concurrently, dominant fish reduce costs of resource depletion by displacing subordinate group members from good food patches. Dominance interactions within a group reduce the amount of time subordinates spend feeding (subordinate individuals fed at higher rates following the removal of a dominant) and limit a subordinate's access to high quality resources. This combination of benefits and reduced costs ultimately contributes to the economic defensibility of a striped parrotfish territory and has led to the evolution of group territorial behavior in the absence of kin selection and cooperative parental care.     

Tug-of-war over reproduction in a cooperatively breeding cichlid

In group-living animals, dominants may suppress subordinate reproduction directly and indirectly, thereby skewing reproduction in their favour. In this study, we show experimentally that this ability (‘power’) is influenced by resource distribution and the body size difference between unrelated dominants and subordinates in the cichlid Neolamprologus pulcher. Reproduction was strongly skewed towards the dominant female, due to these females producing more and larger clutches and those clutches surviving egg eating better than those of subordinate females, but was not so when subordinates defended a patch. If breeding shelters were provided in two patches, subordinate females were more likely to exclusively defend a patch against the dominant female and breed, compared to when the same breeding resource was provided in one patch. Relatively large subordinate females were more likely to defend a patch and reproduce. Females also directly interfered with each other’s reproduction by eating the competitors’ eggs, at which dominants were more successful. Although dominant females benefited from subordinate females due to alloparental care and an increase in egg mass, they also showed costs due to reduced growth in the presence of subordinates. The results support the view that the dominant’s power to control subordinate reproduction determines reproductive partitioning, in agreement with the predictions from tug-of-war models of reproductive skew.     

Dominance status influences female reproductive strategy in a territorial African rodent <Emphasis Type="Italic">Rhabdomys pumilio</Emphasis>

Dominance status influences the fitness of many mammals. Using African striped mice Rhabdomys pumilio, we tested whether (1) dominant females have greater reproductive success than subordinate females, (2) dominant females influence the reproductive output of subordinate females when they are housed in close proximity, (3) reproductive output of a female changes in response to the dominance status of her neighbours, and (4) whether prolonged association between individuals influences the variance in reproductive success between dominants and subordinates (i.e. the ‘dear enemy’ phenomenon). The size and mass of litters of dominants increased significantly when housed adjacent to subordinates than when housed apart. The litter size and mass of subordinates remained unchanged, although subordinates spent significantly more time with their pups when housed close to dominants than when housed apart; time spent with pups by dominants remained unchanged. Moreover, females modified their reproductive output and behaviour in relation to the dominance status of their neighbours. Following prolonged association, dominants still had greater reproductive success, but now, the time spent with pups decreased in subordinates. We suggest that dominants adopt a strategy to increase the reproductive value of their litter, whereas subordinates adopt a pup defence strategy. These strategies are flexible and are influenced by the dominance status and period of association between neighbours, so that females could maximize their fitness in response to varying social conditions.     

Length of feeding day and body weight of great tits in a single- and a two-predator environment

The risk of predation may influence the acquisition of energy and the feeding activity of animals. Feeding activity and body reserves of wintering great tits Parus major in response to the priority to food access were studied in two areas differing in incidence of predators. The one-predator area contained sparrowhawks Accipiter nisus only, whereas the two-predator area contained both sparrowhawks and pygmy owls Glaucidium passerinum, whose hunting periods overlap at dawn and dusk. In the two-predator area dominant great tits arrived at feeders significantly later in the morning, and left earlier in the evening, than their subordinate flock-mates. Hence, feeding day length of dominants was found to be significantly shorter. The reverse was true for the one-predator area. In addition, dominants carried significantly greater reserves than subordinates in the area inhabited by two predators. Factors constraining subcutaneous energy reserves were also studied in removal experiments. After the removal of dominant individuals, subordinate great tits did not reduce their body reserves in the two predator area. In contrast, subordinate great tits significantly reduced evening body reserves in the single-predator area. I concluded that the presence of the two predators increases unpredictability in feeding conditions for great tits. Dominant individuals responded to this by shortening their feeding day and increasing body reserves at dusk. Received: 8 December 1999 / Received in revised form: 15 March 2000 / Accepted: 31 March 2000     

Social dominance and migratory restlessness in the dark-eyed junco (Junco hyemalis)

Summary I investigated the effect of restricted food and social dominance on nocturnal migratory activity (Zugunruhe) in dark-eyed juncos (Junco hyemalis) in late fall and winter. Highly restricted food tended to increase Zugunruhe in both dominant and subordinate members of pairs, however, subordinates showed significantly more migratory activity than dominants or solitary controls. Further, subordinate birds continued Zugunruhe after dominants and solitary controls had ceased this activity for the remainder of the winter. From mid-December through mid-January when birds had access to food ad lib, migratory activity decreased significantly. These results indicate that toward the end of fall migration, migratory behavior is subject to ecological and social conditions that influence the probability of overwinter survival. Presumably, such a system enables these birds to minimize the distance of migration; at the same time, they are able to track a relatively unpredictable, temperate, winter environment by prolonging migration if necessary.     

Reproductive suppression and inbreeding avoidance in wild populations of co-operatively breeding meerkats (Suricata suricatta)

Meerkats live in co-operatively breeding familial groups in which reproduction is monopolised by a dominant pair of breeders. Offspring of the breeders are behaviourally subordinate, and typically remain in their natal group as sexually mature, non-breeding helpers. In this study, we investigated the proximate factors limiting subordinate reproduction. Evidence for reproductive suppression by dominants was investigated by comparing life history, behaviour and hormonal profiles of dominants and subordinates. Baseline levels of plasma luteinising hormone (LH) were significantly higher in dominant than in subordinate females. However, following an exogenous injection of gonadotrophin-releasing hormone (GnRH), both categories had comparable concentrations of circulating LH. There were no significant differences in pre- or post-GnRH challenge LH levels in dominant or subordinate males. Reproduction in both dominant and subordinate females rarely occurred in the absence of unrelated males. Given that groups typically comprise parents and offspring, lack of suitable mates emerged as the primary constraint on subordinate reproduction. When this constraint was removed, subordinates typically bred but at a lower rate than dominants. This difference in reproduction may be attributed to intrasexual competition manifested through direct interference by dominant females through subordinate evictions, infanticide and the abandoning of subordinate litters. We argue that differences in reproductive regulation within mammalian co-operative breeding systems may be explained by differences in the mating strategy (inbreeding versus outbreeding) and the probability that subordinates in obligate outbreeding species will encounter unrelated opposite-sex partners. Received: 19 April 2000 / Accepted: 17 July 2000     

Subordinate males are more likely to help if unrelated to the breeding female in cooperatively breeding white-browed scrubwrens

Subordinates in communally breeding groups of birds usually help to provision nestlings, but in some species helping-at-the-nest is facultative. In species in which groups usually contain relatives, subordinates either always feed young or are more likely to do so when breeding dominants are close relatives, suggesting that benefits of helping collateral kin are important. In other species, adult group members are unrelated to each other and males may only feed young if they have gained paternity, showing that cooperation is related to the mating system. The white-browed scrubwren, Sericornis frontalis, is a communally breeding species in which most groups consist of a simple pair or a dominant pair with a subordinate male. Subordinate males either fed nestlings in a given nest at a rate comparable to the dominants, or did not feed them at all. Breeding groups usually formed through natal philopatry of males, so that about 80% of subordinates were closely related to one or both members of the dominant pair. However, because of death and dispersal, 54% of subordinates were unrelated to the resident female. Although subordinates with their mother fed nestlings in 48% of cases, they fed offspring in 75% of cases if their mother had been replaced by an unrelated female, suggesting that their decision to help is influenced by the opportunity to mate with the female. Supporting this conclusion, relatedness to the dominant male did not affect subordinate behaviour, and genetic studies showed that subordinates often gained paternity if unrelated to the female. Thus, paradoxically, provisioning nestlings is related to the opportunity for mating in a society in which there is natal philopatry and subordinates are usually related to one or both members of the dominant pair. Received: 25 January 1997 / Accepted after revision: 1 May 1997     

Rank differences in energy intake rates in white-faced capuchin monkeys, Cebus capucinus: the effects of contest competition

The effect of aggressive competition over food resources on energy intake rate is analyzed for individuals of three groups of 25–35 white-faced capuchin monkeys, Cebus capucinus, living in and near Lomas Barbudal Biological Reserve, Costa Rica. An individuals energy intake rate on a given food species was affected by its rank and the number of agonistic interactions within the feeding tree. Dominant group members had higher energy intake rates relative to subordinate group members whether or not there was agonism within the feeding tree. Low- and mid-ranked individuals had lower energy intake rates in trees with higher amounts of aggression, while energy intake rate of high-ranked individuals was not affected by the amount of aggression in the feeding tree. Energy intake was not influenced by the sex of the individual when rank was held constant statistically. Energy intake was positively correlated with total crown energy (measured in kilojoules) within the feeding tree for two of three study groups. This difference may be explained by the quality of each groups territory. Finally, high-ranked individuals are responsible for the majority of agonism within feeding trees and target middle- and low-ranked individuals equally. These findings fit the predictions of current socioecological models for within-group contest competition over food resources. The results of this study suggest that within-group competition affects energy intake rate in white-faced capuchin monkeys.     

Reproductive sharing and proximate factors mediating cooperative breeding in the African wild dog (Lycaon pictus)

Although dominant African wild dogs (Lycaon pictus) are generally believed to be the sole breeders within a pack, earlier behavioral and endocrine data suggest that reproduction could be shared with subordinates. We performed an extensive behavioral, demographic, and genetic evaluation of a wild dog population in South Africa to examine the level of such sharing and the proximate mechanisms influencing reproductive contributions of each sex. While a majority of pups were born to dominants because of a lack of subordinate potential breeders, we discovered a substantial portion of reproductive sharing between dominants and subordinates. Compared with alpha females that mated annually, subordinate beta females bred in 54.5% of years whereas thetas never bred. The three top-ranking males all sired pups (56.0%, 32.0%, and 12.0%, respectively) when three or more adult males were present. With only two pack males, alpha and beta individuals shared reproduction nearly equally (55.2% and 44.8%, respectively), and litters of mixed paternity were discovered on eight of 15 (53.3%) occasions. A skewed adult sex-ratio and frequent alpha mortalities for females and behavioral aggression in males allowed most individuals to attain dominant status in their lifetime, creating a constantly shifting social hierarchy. Genetic parentage results corresponded to reported hormone profiles, suggesting physiological suppression in some lower-ranked individuals of both sexes. Thus, a combination of demographic, behavioral, and hormonal proximate factors mediates reproductive partitioning in wild dogs. We conclude that reproductive sharing can be significant in this species, especially for males that have less robust suppressive mechanisms than females.     

Evidence of kin-selected tolerance by nestlings in a siblicidal bird

Behaviorally dominant members of blue-footed booby (Sula nebouxii) broods can effect siblicide by restricting access of subordinate siblings to parents providing food. In spite of their capacity for siblicide, dominant chicks permit subordinates to feed during short-term food shortage; in fact, the proportion of the food that the dominant takes is independent of the total amount delivered in older chicks. A model of optimal food distribution suggests that dominant chicks maximize their inclusive fitness with this pattern, rather than by satisfying their own food requirements and leaving what remains for the subordinate sibling. The indirect reproductive potential represented by a chick's sibling appears to have influenced the evolution of siblicidal brood reduction in this species.     

Aggression and submission reflect reproductive conflict between females in cooperatively breeding meerkats Suricata suricatta

In many cooperatively breeding species, dominant females suppress reproduction in subordinates. Although it is commonly assumed that aggression from dominant females plays a role in reproductive suppression, little is known about the distribution of aggressive interactions. Here, we investigate the distribution of aggressive and submissive interactions among female meerkats (Suricata suricatta). In this species, dominant females produce more than 80% of the litters, but older subordinates occasionally breed. Dominant females commonly kill the pups of subordinates and usually evict older female subordinates from the group 1–3 weeks before the birth of the dominant female's litter. The aggression frequency of the dominant female toward subordinates and the submission frequency that each subordinate female showed to the dominant female increased as the age of the subordinate female increased and as the birth of the dominant female's pups approached. Moreover, as birth approached, both of these behaviors intensified more quickly between the dominant female and older subordinates than between the dominant female and younger subordinates. The aggression frequency of the dominant female toward each subordinate female predicted whether that subordinate female was evicted from the group; the submission frequency by each subordinate female predicted the timing of their eviction during the pregnancy period of the dominant female. These results support the idea that conflict between dominant and subordinate females increases with the age of subordinates and, since older subordinate females are most likely to reproduce, suggest that dominant females may less easily control reproductive attempts by older subordinate females.     

Seasonal variation in feeding behavior, competition and female social relationships in a forest dwelling guenon, the blue monkey (Cercopithecus mitis stuhlmanni), in the Kakamega Forest, Kenya

Socioecological models relate differences in feeding strategies to variation in the nature of female social relationships. Among the African forest guenons, females consume large quantities of fruit and other plant reproductive parts, resources which are thought to promote contest competition, yet these monkeys have been characterized as having agonistically undifferentiated relationships in which rank, if discernible at all, does not correlate with fitness benefits. To determine whether female relationships become more hierarchical under relevant ecological conditions, we monitored the adult females of two blue monkey groups (Cercopithecus mitis stuhlmanni) over a complete annual cycle in the Kakamega Forest, Kenya. Females competed aggressively for plant reproductive parts more often than any other resource type, and in both groups we detected linear dominance hierarchies. Nonetheless, agonism rates remained low throughout our study, and did not vary with changes in ecological conditions. Rather, when plant reproductive parts were scarce, subordinate females spent more time feeding and less time resting in an apparent attempt to compensate for a reduced efficiency of food intake. The effects of rank and food abundance were not reflected, however, in the distribution of grooming. The use of alternative feeding strategies appeared to blunt competition – females of all ranks were unlikely to be near others while feeding and spent more time consuming alternative resources when plant reproductive parts were scarce. The diverse diet of this species may allow females to avoid conflict so that dominance has only subtle effects that are difficult to detect. While socioecological models often simplify the connection between resources and female interactions, our results emphasize that the behavior of animals consuming particular resources, and not the resources themselves, are critical predictors of social patterns.     

Social and ecological influences on activity budgets of vervet monkeys,and their implications for group living

Summary Activity patterns were documented over a 20-month period in six groups of vervet monkeys (Cercopithecus aethiops) in Amboseli National Park, Kenya. Group size varied both among groups and within groups through time. The home ranges of two groups were located in Acacia xanthophloea habitat whereas those of four adjacent groups were located in A. tortilis habitat. Repeated measures analysis of variance, three-way analysis of variance, and meta-analysis of group size effects were carried out on feeding, moving, resting, scanning, allo-grooming, and other behavior. Time spent feeding varied seasonally, peaking during periods when A. tortilis seeds were eaten. However, time spent feeding did not vary within or among groups, or between habitats, males and females, or dominants and subordinates. Males spend more time scanning and less time allo-grooming than females, and high-ranking individuals of both sexes spent more time scanning than low-ranking individuals. Among females, subordinates spent more time moving, whereas among males, dominants spent more time moving. Groups living in A. xanthophloea habitat spent more time moving and allo-grooming than groups in A. tortilis habitat but this may have been confounded by the fact that these two groups were on average larger than the others. Significant differences in activity budgets among groups demonstrate that activity patterns observed in one group are not always readily generalized to other groups even when they come from the same population. Group size analyses examined potential costs and benefits of group living. Increased intragroup competition, measured by time spent moving and feeding, was only weakly evident in larger groups. Individuals in larger groups allo-groomed more than individuals in smaller groups. Self-grooming also increased in larger groups. Individuals in larger groups spent less time scanning than individuals in smaller groups, but there was more scanning per group in larger groups than in smaller groups. Correspondence to: L.A. Isbell     

Aggresive competition and individual food consumption in wild brown capuchin monkeys (Cebus apella)

Summary The impact of aggresive competition on food intake at all the resources used is analysed for every member of a group of brown capuchin monkeys (Cebus apella) in the Manu National Park, Peru, where they live in groups of 8–14 animals. An individual's food intake at a given tree was affected independently both by its domirance rank (Fig. 1) and by how much aggression it received (Table 5). Food intake was not strongly affected by body size when dominance rank was held constant by partial correlation. At food sources where high rates of fighting occurred, an individual's food intake depended more on its domirance status than on the rate of aggression it received (Fig. 2). However, food intake at resources where rates of fighting were low depended mostly on the rate of aggression received. When aggression over food was absent, the food intakes of dominants and subordinates were equal. Dominants had significantly greater total energy intake (20.5% more) than did subordinates, even though more than one third of their diet came from food sources where little or no fighting occurred (Fig. 3). Energy intake was also significantly greater for individuals that received little aggression. The only adult that emigrated from the main study group was the individua with the lowest energy intake. Competition for food within groups was more than ten times as intense as competition between brown capuchin groups.     

Social foraging and dominance relationships: the effects of socially mediated interference

In socially foraging animals, it is widely acknowledged that the position of an individual within the dominance hierarchy of the group has a large effect upon its foraging behaviour and energetic intake, where the intake of subordinates can be reduced through socially mediated interference. In this paper, we explore the effects of interference upon group dynamics and individual behaviour, using a spatially explicit individual-based model. Each individual follows a simple behavioural rule based upon its energetic reserves and the actions of its neighbours (where the rule is derived from game theory models). We show that dominant individuals should have larger energetic reserves than their subordinates, and the size of this difference increases when either food is scarce, the intensity of interference suffered by the subordinates increases, or the distance over which dominant individuals affect subordinates increases. Unlike previous models, the results presented in this paper about differences in reserves are not based upon prior assumptions of the effects of social hierarchy and energetic reserves upon predation risk, and emerge through nothing more than a reduction in energetic intake by the subordinates when dominants are present. Furthermore, we show that increasing interference intensity, food availability or the distance over which dominants have an effect also causes the difference in movement between ranks to increase (where subordinates move more than dominants), and the distance over which dominants have an effect changes the size of the groups that the different ranks are found in. These results are discussed in relation to previous studies of intra- and interspecific dominance hierarchies.     

The special role of male Cebus monkeys in predation avoidance and its effect on group composition

Summary Vigilance behavior, predator detection abilities, and responses to real and model predators were studied in two species of capuchin monkeys (Cebus albifrons and C. apella) in a Peruvian lowland rain forest. Adult males were more vigilant than adult females in both species, mainly because the males spent less time feeding and foraging and partly because they were at the periphery more often than the females. The increased vigilance of adult males is reflected in their superior performance in the detection of (model) predators. Adult and subadult males were also far more likely to approach and mob real and model predators. Adults that were outside the center of the group increased foraging activities but cut back an feeding, much of which was done in exposed tree crowns. Current theory suggests that primate groups are multi-male when a single male is unable to defend sexual access to the group of females. In these small capuchin groups, which are multimale, the enhanced safety of females and young provided by extra adult males furnishes a more plausible explanation. A comparison of the two capuchins with the ecologically similar Southeast Asian Macaca fascicularis suggests that the high predation risk outside the group may also have caused the unusual male career profile in capuchins, which have a long tenure of dominants and a very long potential lifespan. Further predictions of this hypothesis are developed.     

Why is competition more intense if food is supplied more slowly?

We investigated competition for food among two groups of six clone amagos (salmonids), Oncorhynchusmasoumacrostomus, in a laboratory experiment with different rates of food input. We examined the effect of temporal clumping of food resources on the inequality of food sharing between competitors. Monopolization of food by dominants was greater at a low input rate (one food item per 10 s) than at a high input rate (1 food item per 1 s). Aggressive behavior by dominants was more frequent at the low input rate than at the high input rate; its purpose was presumably to interfere with the feeding behavior of subordinates. We assessed the relative importance of three foraging factors (the number of approaches to food items, the chance per approach and the gain per chance) in enhancing inequality in food gain between individuals. Dominants had a disproportionately high chance per approach and gain per chance at the low input rate, but not at the high input rate. The chance of obtaining a food item per approach depended on how many competitors approached simultaneously. The gain per chance depended on the competitive ability of the approaching fish. There was an interaction between these components, such that the number of approaches affected the chance per approach and gain per chance. We evaluated the independent effect of the chance per approach, and showed that it was higher for dominants than for subordinates at the low input rate, but not at the high input rate. This implies that subordinates changed their behavior and became more likely to avoid approaching food at the same time as dominants at the low input rate. Received: 13 August 1996 / Accepted after revision: 30 November 1996     

Fattening strategies in dominance-structured greenfinch (Carduelis chloris) flocks in winter

Relationships between social status and levels of body reserves stored by members of greenfinch (Carduelis chloris) flocks in winter were investigated. In addition, the adjustment of reserves by birds of different rank to experimental changes in food predictability and to changed weather conditions were examined. Birds with low social status carried overall larger body reserves than high-ranked birds. The results of the experiments suggest that this was mainly because subordinates, due to their low priority to food access, perceived future foraging success as less predictable than dominants. In response to severe weather, which probably increased the risk of starvation for birds, dominants temporarily increased their reserves more than subordinates. This response also indicated that birds with low social status carried larger reserves than high-ranked birds, and demonstrated that dominants could increase food intake when there was a risk of energetic shortfall. The results suggest that fattening strategies in greenfinches depend on social status. In winter, subordinates may be forced to carry larger reserves than dominants to safeguard against social constraints in access to food during critical times. As body reserves may be costly to carry and acquire, this should reduce the probability of surviving the winter for subordinates compared to dominants. Received: 2 March 1995/Accepted after revision: 13 April 1996     

Nonlinear effects of food aggregation on interference competition in mallards

Previous studies of interference competition have shown an asymmetric effect on intake rate of foragers on clumped resources, with only subordinate individuals suffering. However, the food distributions in these studies were uniform or highly clumped, whereas in many field situations, food aggregation is intermediate. Here we investigated whether food distribution (i.e., uniform, slightly clumped, and highly clumped) affects the behavioral response of mallards foraging alone or competing with another. Although the amount of food was the same in all distributions, the mallards reached higher intake rates, visited fewer patches, and showed longer average feeding times in the highly clumped distribution. Competing mallards had lower intake rates on the slightly clumped than on the uniform or highly clumped food distributions. Subordinates generally visited more patches and had shorter feeding times per patch, but their intake rates were not significantly lower than those of dominants. Therefore, we propose that subordinates do not necessarily suffer from interference competition in terms of intake rate, but do suffer higher search costs. In addition, although dominants had significantly higher average feeding times on the best quality patches of the highly clumped food distribution, such an effect was not found in the slightly clumped distribution. These findings indicate that in environments where food is aggregated to a lesser extent, monopolization is not the best strategy for dominants. Our results suggest that interference experiments should use food distributions that resemble the natural situation animals are faced with in the field.