Use of different signaling modalities to communicate status by dominant and subordinate Heermann’s kangaroo rats (<Emphasis Type="Italic">Dipodomys heermanni</Emphasis>)

Dominance relationships in solitary species may be an important factor in the maintenance of long-term, stable relationships among territorial neighbors. We examined the mediation of intraspecific interactions in a solitary, territorial kangaroo rat, Dipodomys heermanni, and tested whether unfamiliar kangaroo rats establish a dominance hierarchy and then decrease aggression and increase communication (via footdrumming and sandbathing) after initial interactions and the establishment of a social structure. Results revealed that both dominance hierarchies and familiarization with particular individuals are likely to mediate social interactions. After only one pairing per dyad, an almost linear dominance hierarchy emerged, which became perfectly linear after a 90-min familiarization period. During the course of subsequent interactions between dyad partners, fighting decreased and non-agonistic communication increased. Dominant kangaroo rats sandbathed at higher rates than subordinates, possibly to deposit scent to advertise competitive ability, whereas subordinate kangaroo rats footdrummed from inside the burrow, which seemed to indicate an unwillingness to interact. We suggest the kangaroo rats use a conditional strategy when deciding to fight (be dominant) or withdraw (be subordinate) by employing different modes to communicate status and minimize the potential risk of injury during unnecessarily prolonged fights.     

Queuing for dominance: gerontocracy and queue-jumping in the hover wasp <Emphasis Type="Italic">Liostenogaster flavolineata</Emphasis>

The mechanisms through which dominance is inherited within social groups vary from direct interactions such as fighting to non-confrontational conventions. Liostenogaster flavolineata is a primitively eusocial hover wasp in which one female, the ‘dominant’, is the only reproductive upon the nest. The remaining females, although capable of reproduction, behave as helpers. In this study, we investigate the rules by which helpers inherit dominance. We removed successive dominants from 56 nests and recorded accession on un-manipulated nests. The results showed that L. flavolineata has a strict age-based inheritance queue: new dominants are the oldest female in their groups 87% of the time. Thirteen cases of queue-jumping were found in which young individuals were able to supplant older nestmates and inherit dominance precociously. Queue jumpers did not differ from other wasps in terms of relatedness to other group members or body size. Individuals that had previously worked less hard than other females of equivalent rank were significantly more likely to later jump the queue. Queue-jumping may represent a cheating strategy or could indicate that the rule for inheriting dominance is not based purely on relative age. We also discuss possible reasons why age-based queuing has evolved and its potential to promote the evolution of helping behaviour.     

Who’s what? Prompt recognition of social status in crayfish

Crayfish are excellent model organisms to study the proximate mechanisms underlying the maintenance of dominance hierarchies in invertebrates. Our aim here was to investigate whether Procambarus clarkii males use social eavesdropping to discriminate dominant from subordinate crayfish. To this end, we conducted an experiment composed of a “passive” and an “active” phase. In the passive phase, “focal” individuals were allowed (treatment 1) or not (treatment 2) to see and smell two size-matched crayfish fighting while, in the subsequent active phase, they were allowed to freely interact with the fighting dyad. None of the recorded variables showed any significant difference between the two treatments, but, invariably, focal individuals were able to promptly discriminate dominant from subordinate crayfish. This study provides evidence that male crayfish recognize the social status of a conspecific without the need of direct or indirect experience with it and avoid dominants—and thus dangerous opponents—by means of a badge of status. A form of “winner and loser effects” could also contribute to the structuring of dominance/subordinate relationships. The implication of these results in understanding the maintenance of dominance hierarchies in invertebrates are discussed and compared with findings previously achieved in the context of mate choice by P. clarkii females, who do appear to use eavesdropping to identify dominants and subordinates.     

Fighting rules and rival recognition reduce costs of aggression in male lizards, Podarcis hispanica

The establishment of fighting rules and the ability to recognise individual conspecifics and to assess their fighting ability and/or roles may help to reduce costs of fighting. We staged encounters between males of the lizard Podarcis hispanica to examine whether lizards used fighting strategies and whether a previous agonistic experience affects the outcome and characteristics of a subsequent encounter. The results showed that simple rules such as body size differences and residence condition were used to determine the outcome of agonistic interactions as quickly as possible. Thus, larger males were dominant in most encounters. However, when size differences between opponents are smaller, they may be more difficult to estimate and, then, residence condition was more important. In addition, the intensity of interactions between males could be explained according to the ”sequential assessment game”, supporting the idea that P. hispanica males acquire information about fighting ability gradually during the progress of a fight. Our results also showed that the second fight of the same pair of males was less aggressive, even when its outcome was the opposite of the first. This result suggests that male P. hispanica can recognise individual opponents and that they use this information to reach a contest outcome more quickly, thus reducing unnecessary aggression levels in subsequent interactions. These fighting strategies and assessment mechanisms may help to stabilise the social system of this lizard. Received: 2 November 1999 / Revised: 26 August 2000 / Accepted: 4 September 2000     

A game model for dominance relations among group-living animals

We present here an attempt to understand behaviors of dominant individuals and of subordinate individuals as behavior strategies in an asymmetric “hawk-dove” game. We assume that contestants have perfect information about relative fighting ability and the value of the resource. Any type of asymmetry, both relevant to and irrelevant to the fighting ability, can be considered. It is concluded that evolutionarily stable strategies (ESSs) depend on the resource value (V), the cost of injury (D), and the probability that the individual in one role will win (x). Different ESSs can exist even when values of V, D, and x are the same. The characteristics of dominance relations detected by observers may result from the ESSs that the individuals are adopting. The model explains some characteristics of dominance relations, for example, the consistent outcome of contests, the rare occurrence of escalated fights, and the discrepancy between resource holding potential (RHP) and dominance relations, from the viewpoint of individual selection. Received: 7 May 1997 / Accepted after revision: 17 October 1997     

Prior residency as a key determinant of social dominance in the willow tit (Parus montanus)

Summary We studied how age, body size and prior residency affected social dominance in the willow tit (Parus montanus) groups. The contribution of each variable was experimentally tested in unisexual two-bird trials, in which the birds were matched for all variables except the one studied. Large birds were dominant over smaller ones (Fig. 1). The effect of body size was more prominent in males than in females. Age had no influence on dominance. Residents became dominant more often than newcomers (Fig. 2). Adulthood or larger body size did not override the advantage of prior residency (Fig. 2). Therefore, the proximate reason for the age-dependent dominance seen in natural willow tit flocks is most likely the prior residency advantage of the adults. Factors connected with fighting ability (body size and age) seem to be less important than the time of establishment of rank, which may reflect the importance of resource value differences between residents and newcomers in this context. The advantage of residency might make it advantageous to be a member of a flock even as a subordinate, rather than being solitary.     

Correlations of food distribution and patch size with agonistic interactions in female vervets (Chlorocebus aethiops) and patas monkeys (Erythrocebus patas) living in simple habitats

Food distribution is hypothesized to be important in determining the nature of female relationships within social groups of primates. When food limits female reproductive success, spatially clumped foods are expected to produce strong, linear dominance hierarchies within groups, whereas more spatially dispersed foods are expected to produce weaker or non-existent dominance hierarchies. The association between food distribution and competitive relationships presumably occurs because clumped foods are usurpable but dispersed foods are not. We examined the spatial distribution of food patches (trees) and patch size relative to feeding behavior and agonistic interactions in vervets and patas monkeys, two closely related and sympatric species that nonetheless differ in the strength of the female dominance hierarchy. Food patches of both patas monkeys and vervets were small in size and randomly distributed in Acacia drepanolobium habitat. In contrast, in A. xanthophloea woodland, the habitat type that was exclusively used by vervets, food patches were larger and more spatially clumped. These similarities and differences between and within species were correlated with similarities and differences in the strength and linearity of their dominance hierarchies. Patas monkeys and vervets in A. drepanolobium habitat had dominance hierarchies that were weakly defined because there were relatively few agonistic interactions between females. By contrast, in A. xanthophloea habitat, vervets had a stronger, linear dominance hierarchy characterized by a higher rate of agonistic interactions over food. The covariation of agonistic interactions with patch size is discussed in relation to depletion time, another characteristic that may covary with food distribution, and resource renewal rate, an important determinant of agonistic interactions in insectivorous birds, fishes, insects, and mammals. Received: 18 February 2000 / Revised: 5 September 2000 / Accepted: 26 September 2000     

Competitive regimes in forest-dwelling Hanuman langur females (Semnopithecus entellus)

Van Schaik’s socioecological model predicts interrelations among food distribution, competitive regimes, and female social relationships. To test the internal consistency of the model, feeding competition was examined in three differently sized groups of a forest-dwelling population of Hanuman langurs (Semnopithecus entellus). The nutritional condition of females was used as a direct indicator of feeding competition and related to the seasonal variation in resource distribution and abundance. Female dominance hierarchies were characterized by displacements. Dominance hierarchies were significantly linear and relatively stable, but less so with increasing group size. Physical condition correlated with dominance rank and high-ranking females were in the best condition, indicating within-group contest competition. The strength of this relationship became less pronounced with increasing group size. The females of the medium-sized group were in the best physical condition indicating between-group contest plus within-group scramble competition. Closer examination revealed variable costs and benefits of group foraging with a predominance of within-group scramble competition when food was more abundant. The results support some basic predictions of the model. Limiting food abundance was bound to ubiquitous within-group scramble competition. The use of clumped resources translated into differences in net energy gain based on dominance. In contrast to the predictions, group-size-related costs and benefits were related to food abundance instead of food distribution. As predicted, within-group contest competition was linked to a linear dominance hierarchy. The absence of nepotism and coalitions in Hanuman langurs may be attributed to dominance hierarchies that are unstable through time, probably minimizing fitness gain via kin support. Received: 25 May 1999 / Received in revised form: 18 February 2000 / Accepted: 25 February 2000     

Fighting behavior in hermit crabs: the combined effect of resource-holding potential and resource value in Pagurus longicarpus

A study was conducted to determine whether asymmetries in both resource-holding potential (RHP) and resource value (RV) influence dominance and fighting behavior in the hermit crab Pagurus longicarpus. A total of 120 groups of three crabs were observed for 10 min in four experiments that tested all diverse combinations of equal/different RHP (i.e. ‘body size’) and equal/different RV (i.e. ‘shell size’ and ‘shell quality’). In a fifth experiment, dominant and subordinate individuals of the same size category (26 groups) were forced to enter shells of opposite quality than those previously occupied, and then the behavior of the reconstituted original groups was observed for additional 10 min. As expected, crabs in lower quality shells were more willing to initiate and to escalate fights. However, their attacks were directed to any crab of the group, independently of the defender's shell quality, and the fight duration did not vary with the different value of the resources at stake. This may indicate that P. longicarpus is unable to assess the quality of the shells available in its social environment but bases its tactical decisions during fights solely on its own resource. This suggestion was confirmed by the change in the fighting behavior of crabs whose shell quality was experimentally altered. This manipulation induced an overall increase in the intensity of aggression, drastic modification of crab behavior, and inversion of the hierarchy even though these crabs have had previous experiences of wins/losses and were familiar to the other members of the group. In this species, large crab size and/or the occupancy of adequate (and oversized) shells appeared to be the most likely determinant of contest resolution. Individuals seemed to retain a memory of the previously held resource and behaved accordingly.     

Dominance and reproduction in a parthenogenetic lizard

Summary In most vertebrates, males and females are believed to differ in terms of their investment in offspring. Dominance theory suggests that one way individuals of the sex with lower parental investment can increase reproductive success would be to dominate others of the same sex. The dominant competitors are thought to achieve preferred access to mates, and thus, have greater reproductive success than subordinates.Reproduction in parthenogenetic Cnemidophorus uniparens normally proceeds without males, but individuals exhibit mounting behaviors in captivity that are typical of closely related Cnemidophorus species that reproduce sexually. Thus, these animals provide an unusual opportunity to study the effects of behavior on reproduction apart from copulation and fertilization. In this study relationships between dominance and reproduction were investigated in the unisexual lizard species, C. uniparens. Dominance hierarchies were rapidly established and maintained in the laboratory by agonistic encounters among individuals. The number of times an individual charged its cagemates was positively correlated with the number of clutches and eggs laid. Also, dominant animals who charged their cagemates were likely to win agonistic encounters; recipients of charges usually fled. Hierarchies based on different behaviors were not all related to reproduction. Charges as a predictor of dominance was unrelated to body length, percent increase in body length and time spent in the basking site. However, individuals with a high percent increase in body length spent more time basking. This is likely a result of the increased energy demands of growth in addition to reproduction. Physiological stress as measured by plasma corticosterone titers was unrelated to dominance. We suggest that dominance is an important factor affecting reproduction in C. uniparens.     

Lifetime mating success, sexual selection and life history of fallow bucks (Dama dama)

We used data from a long-term study (15 years) of fallow deer to report for the first time the lifetime mating success, overall variance in lifetime mating success, and age-specific mortality levels of males. Fallow bucks that gain matings have higher social dominance rank, higher rates of fighting, and invest more in vocal display during the breeding season than unsuccessful males. Therefore, we examined if mating was associated with trade-offs in terms of survival, lifespan, and mating potential. We found that the variance in lifetime mating success was very high: 34 (10.7%) males mated, and of those, the 10 most successful males gained 73% of all matings (n=934). Mortality rates were generally high and only 22.3% (71/318) of males reached social maturity, i.e., 4 years. The oldest male was 13 years old. We found that fallow bucks that mated were not more likely to die during the following year, did not suffer from a reduction in lifespan, and did not incur lower mating success later in life as a result of mating during the early years of social maturity. Our results show that mating males at age 5 years (and possibly 9 years) may be more likely to survive than non-mating males. Additionally, the number of matings gained by males during the first years of social maturity was positively correlated with lifespan. We suggest that mating males are of higher quality than non-mating males because they are not more likely to incur trade-offs as a result of their increased reproductive efforts. Received: 9 November 1999 / Revised: 30 April 2000 / Accepted: 27 May 2000     

Dominance and polyethism in the eusocial wasp Mischocyttarus mastigophorus (Hymenoptera: Vespidae)

Dominance interactions affected patterns of non-reproductive division of labor (polyethism) in the eusocial wasp Mischocyttarus mastigophorus. Socially dominant individuals foraged for food (nectar and insect prey) at lower rates than subordinate individuals. In contrast, dominant wasps performed most of the foraging for the wood pulp used in nest construction. Social dominance also affected partitioning of materials collected by foragers when they returned to the nest. Wood pulp loads were never shared with nest mates, while food loads, especially insect prey, were often partitioned with other wasps. Dominant individuals on the nest were more likely to take food from arriving foragers than subordinate individuals. The role of dominance interactions in regulating polyethism has evolved in the eusocial paper wasps (Polistinae). Both specialization by foragers and task partitioning have increased from basal genera (independent-founding wasps, including Mischo-cyttarus spp.) to more derived genera (swarm-founding Epiponini). Dominance interactions do not regulate forager specialization or task partitioning in epiponines. I hypothesize that these changes in polyethism were enabled by the evolution of increased colony size in the Epiponini. Received: 8 December 1997 / Accepted after revision: 28 March 1998     

The role of body size and fighting experience in predicting contest behaviour in the black field cricket, <Emphasis Type="Italic">Teleogryllus commodus</Emphasis>

Body size strongly predicts fighting behaviour and outcome in many species, with the larger opponent usually winning contests. However, recent fighting experience can have a strong influence on the establishment of dominance hierarchies, with recent winners being more likely to win subsequent contests, while recent losers are more likely to lose. Recent fighting experience therefore has the potential to modify the effect of body size in determining contest behaviour and outcome. Here, we investigate whether recent fighting experience weakens the role of body size in predicting contest behaviour in the black field cricket, Teleogryllus commodus. We compared the role of body size in determining contest outcome during initial non-physical encounters and escalated aggressive physical encounters (grapples), as well as the probability of escalation occurring, in contests involving either a naïve or experienced smaller male against a naïve larger male. We found that recent fighting experience only affected contest outcome during non-physical encounters. Once a contest had escalated into grappling, the effect of previous wins and losses was no longer apparent and body size strongly predicted contest outcome. Thus, once males can directly assess their opponent's fighting ability, recent fighting experience did not alter the effect of body size on contest behaviour and outcome.     

Dominance orders in the ponerine ant Pachycondyla apicalis (Hymenoptera,Formicidae)

Summary Intracolony aggression among workers of the neotropical ponerine ant species Pachycondyla apicalis leads to dominance orders. Antagonistic interactions can entail either overt physical attacks with the subordinate individual often exhibiting a submissive posture or the robbing and destruction of eggs laid by nestmates. The single queen, however, was never observed either attacking or being attacked by any colony member. The hierarchical structure among workers consists of one dominant individual and several subordinates; the relationships among subordinate workers are unclear, however. We report for the first time a natural (nonmanipulated) change in the social status of individuals within an ant dominance order. Dominant workers usually had better developed ovaries, laid more eggs and were more frequently observed attending the egg pile than subordinate individuals. This pattern became even more striking when the queen was excluded from the colony. These results indicate that workers of P. apicalis lay eggs even in the presence of the queen. It is possible that some of these haploid eggs may develop into males.Offprint requests to: B. Holldobler     

Behavioral and physiological factors associated with juvenile hormone in Polistes wasp foundresses

Although there is increasing interest in the evolution of endocrine systems, relatively little is known about the factors associated with natural endocrine variation in invertebrates. Here, we assess juvenile hormone (JH) titers among nest-founding queens of the wasp Polistes dominulus over 2 years. We allowed unfamiliar wasps to battle for dominance and examined the relationships between dominance rank, JH, ovarian development, and facial patterns. The relationship between JH-titer and dominance varied across years; there was a stronger relationship between JH-titer and dominance in 2006 than in 2008. Across years, wasps that won dominance contests had facial patterns with more broken black spots than wasps that lost dominance contests. There was no relationship between dominance rank and ovarian development. The individual characteristics associated with JH-titer were also tested; JH-titers were correlated with facial pattern brokenness and ovarian development. This study adds to previous work indicating that P. dominulus facial patterns function as a signal of fighting ability. Furthermore, the correlation between JH-titers and facial patterns parallels previous work on testosterone and vertebrate signals and suggests that links between signals of fighting ability and hormones that mediate fighting ability may be common across taxa. Overall, individual JH-titers vary, though they are typically associated with factors related to individual reproductive success, including dominance, fertility, and facial pattern brokenness. Future studies in additional contexts and taxa will be important to test how and why JH-titers vary.     

Effect of losing on male fights of broad-horned flour beetle, Gnatocerus cornutus

Experience-dependent tactics of males trying to gain access to females were examined in the beetle Gnatocerus cornutus, which engages in male–male fighting for mates. In male fights, subsequent behavior is modified by winning and losing experiences. Males, therefore, may choose between several behavioral tactics to obtain a mate, based on his previous fighting experience. In G. cornutus, we examined for how long aggressive behaviors were modified by experiences of winning or losing, i.e., the duration of the prior experience effect. Losing decreased a male's frequency of fighting for 4 days, and few defeated males fought any male regardless of the opponent's size. By the fifth day, this effect disappeared. No modulation of male behavior due to winning was observed. Furthermore, the experience of losing not only decreased a male's aggressiveness but also switched the male behavior from fighting to dispersal from the fight site to another site. In future, it is necessary to clarify why the optimal term of the losing experience is 4 days in this beetle.     

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.     

Molecular genetic and behavioral analysis of social organization in the Asian elephant (Elephas maximus)

We report on the genetic evaluation and behavioral study of social organization in the Asian elephant (Elephas maximus). Although Asian elephants and African elephants (Loxodonta africana) were previously thought to have similar social organizations, our results demonstrate a substantial difference in the complexity and structure of Asian elephant social groupings from that described for African savanna elephants. Photographic cataloging of individuals, radio telemetry, and behavioral observations in Ruhuna National Park, Sri Lanka, enabled us to assign associated females and young to four groups with overlapping ranges. Genetic sampling of individuals from the four groups in Ruhuna National Park and three other groups in surrounding areas, conducted through PCR amplification and sequencing of mitochondrial DNA from dung, supported the matriarchal nature of female groups and the lack of inter-group transfer of females. Behaviorally and genetically, the identified social groups were best described as ”family groups”. We did not find any evidence for the existence of social groups of higher complexity than family groups. Received: 25 March 2000 / Received in revised form: 28 March 2000 / Accepted: 1 April 2000     

Dominance and survival of dippers Cinclus cinclus

Survival of dippers Cinclus cinclus in the wild was studied in relation to their dominance. Dominance was assessed amongst groups of temporary captives in a laboratory arena. Adults tended to be more dominant than juveniles, and within age classes males generally dominated females. Dominant individuals also sang more. Logistic regression was used to examine survival in the wild in relation to dominance and other factors. Annual survival of juvenile males was negatively related to dominance during the previous autumn. No significant effects of dominance on winter or annual survival were found amongst females or adult males. Dominance effects on overwinter survival approached significance for adult females, however, and were significant for both winter and annual survival when the data from adult females and juvenile males were combined. None of the other factors considered, which included body size, song frequency and year, had a significant effect on overwinter or annual survival. We concluded that dominance either had no influence on survival, or had a negative effect particularly on dippers of intermediate status. We were unable to show if dominance had a direct effect, however, or acted indirectly on survival via an unidentified factor we did not examine. Received: 23 June 1994/Accepted after revision: 28 October 1995     

The social network structure of a wild meerkat population: 2. Intragroup interactions

Knowledge of the structure of networks of social interactions is important for understanding the evolution of cooperation, transmission of disease, and patterns of social learning, yet little is known of how environmental, ecological, or behavioural factors relate to such structures within groups. We observed grooming, dominance, and foraging competition interactions in eight groups of wild meerkats (Suricata suricatta) and constructed interaction networks for each behaviour. We investigated relationships between networks for different social interactions and explored how group attributes (size and sex ratio), individual attributes (tenure of dominants), and ecological factors (ectoparasite load) are related to variation in network structure. Network structures varied within a group according to interaction type. Further, network structure varied predictably with group attributes, individual attributes, and ecological factors. Networks became less dense as group size increased suggesting that individuals were limited in their number of partners. Groups with more established dominant females were more egalitarian in their grooming and foraging competition interactions, but more despotic in their dominance interactions. The distribution of individuals receiving grooming became more skewed at higher parasite loads, but more equitable at low parasite loads. We conclude that the pattern of interactions between members of meerkat groups is not consistent between groups but instead depends on general attributes of the group, the influence of specific individuals within the group, and ecological factors acting on group members. We suggest that the variation observed in interaction patterns between members of meerkat groups may have fitness consequences both for individual group members and the group itself.