Group size, queuing and helping decisions in facultatively eusocial hover wasps

In facultatively eusocial hover wasps, some females leave their natal nests while others choose to stay and become helpers. We tested whether the decisions of 126 newly emerged females to stay or leave depended on group size: the number of females already resident on their natal nests. We predicted that females would be less likely to stay in larger groups, where the benefits of helping are probably smaller and there is a smaller chance of inheriting the dominant position. We also predicted that unrelated females would be less likely to join larger groups. We manipulated group size by removing residents from nests. Newly emerged females disappeared from their natal nests at a rate of 2.5% per day, but did not disappear from manipulated nests at higher rates than controls. Experimentally reducing group size also did not increase the frequency of joiners. Newly emerged females disappeared at twice the rate of older subordinate females, suggesting the existence of a `leaving window' early in life. One problem is the difficulty of distinguishing between leaving and death. Received: 7 July 1998 / Accepted after revision: 25 October 1998     

Social dominance in migrant and resident turkey vultures at carcasses: evidence for a despotic distribution?

We placed carcasses in three different vegetation types in the heterogeneous savannas of central Venezuela to investigate the role of social dominance in habitat use by flocking migrant and resident turkey vultures (Cathartes aura meridionalis and C. a. ruficollis). Migrants foraged primarily in savanna habitats while residents foraged almost exclusively in gallery forest. In the gallery forest residents discovered carrion first significantly more often than migrants, despite there being equal densities of residents and migrants foraging over this habitat. Because residents fed in smaller groups than migrants at carcasses they had higher feeding rates. There was also a negative relationship between group sizes of residents and migrants. The feeding rate of residents declined in response to increased group size of migrants, but group size of residents had no effect. Migrant group size also had a greater effect on resident feeding rates than king vulture presence or absence. When the effect of migrant and resident group size on feeding rates in migrants was compared, the most significant factor was migrant group size. A second analysis showed that both resident group size and presence or absence of king vultures had a significant effect on feeding rates in migrants. Rates of agonistic encounters in migrant and resident turkey vultures increased weakly in relation to group size. However, there was an increase in residents' encounter rate with migrants in relation to increased migrant group size; there was no difference in resident encounter rates with other residents in relation to resident group size. Migrants dominated residents in almost all agonistic interactions over carcasses. We suggest that savanna habitats were less attractive to residents for foraging because they held larger groups of migrants.     

Territory ownership and familiarity status affect how much male root voles (Microtus oeconomus) invest in territory defence

Neighbour–stranger discrimination occurs when individuals respond with more aggression to strangers than to territorial neighbours—a phenomenon termed the “dear enemy phenomenon” (DEP). We investigated the DEP with male and female root voles (Microtus oeconomus Pallas 1776) using field dyadic arena tests conducted in enclosures where we could test for the effects of familiarity (familiar versus stranger), ownership (resident versus intruder status) and resource-holding potential (body mass) on territorial behaviours. The results showed that males put more effort into territorial defence than females, and males could discriminate between neighbours and strangers. In males, aggressiveness was influenced by a significant two-way interaction between treatment and ownership. Male residents were more aggressive towards stranger intruders than towards neighbour intruders, while male intruders were less aggressive towards stranger residents than towards neighbour residents. In females, neither treatment nor ownership status had a significant effect on aggressiveness. Familiar males performed more social behaviours but less non-social behaviours than stranger males. Furthermore, there was a clear dominance hierarchy between residents and intruders in stranger dyads, with the male territory holders dominating the intruder in pairwise interactions. To our knowledge, these results demonstrate for the first time DEP in a small mammal with a known pedigree and present the first evidence for “prior resident advantage” in voles. We argue that both ownership status and familiarity status affect how much an individual invests in territory defence. The benefits of neighbour–stranger discrimination for male root voles and the absence of neighbour–stranger discrimination in female root voles are discussed.     

Shoaling decisions in three-spined sticklebacks (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>)—familiarity,kinship and inbreeding

Shoaling with familiar kin is a well-known phenomenon. It has been described both for adult and for fry of three-spined sticklebacks (Gasterosteus aculeatus). However, evidence of preference for kin independent of familiarity is scarce. Furthermore, inbreeding effects have not been studied for shoaling preferences and there is a lack of studies about changes in individuals’ ability to recognise kin during different phases of life history. We gave inbred and outbred, nonreproductive sticklebacks of different age the choice to shoal with a group of familiar siblings vs unfamiliar non-siblings and with unfamiliar siblings vs unfamiliar non-siblings. Subadult sticklebacks preferred to shoal with familiar kin over unfamiliar non-kin, but inbred and outbred individuals had similar preferences. When given the choice between unfamiliar siblings and unfamiliar non-sibs, adult outbred fish preferred the shoal of kin, while inbred fish behaved indifferently. Body characteristics of the group such as body mass, standard length and condition of its members did not significantly explain the shoaling preferences for kin. Thus, sticklebacks were capable of recognising familiar and unfamiliar kin. The latter capability was lost by inbreeding.     

Parentage and sex allocation in the facultatively polygynous ant Myrmicatahoensis

Most social groups have the potential for reproductive conflict among group members. Within insect societies, reproduction can be divided among multiple fertile individuals, leading to potential conflicts between these individuals over the parentage of sexual offspring. Colonies of the facultatively polygynous ant Myrmicatahoensis contain from one to several mated queens. In this species, female sexuals were produced almost exclusively by one queen. The parentage of male sexuals was more complex. In accordance with predictions based on worker sex-allocation preferences, male-producing colonies tended to have low levels of genetic relatedness (i.e., high queen numbers). Correspondingly, males were often reared from the eggs of two or more queens in the nest. Further, over half of the males produced appeared to be the progeny of fertile workers, not of queens. Overall investment ratios were substantially more male biased than those predicted by genetic relatedness, suggesting hidden costs associated with the production of female sexuals. These costs are likely to include local resource competition among females, most notably when these individuals are adopted by their maternal nest. Received: 3 March 1998 / Accepted after revision: 20 June 1998     

Chemical rival recognition decreases aggression levels in male Iberian wall lizards, <Emphasis Type="Italic">Podarcis hispanica</Emphasis>

In an agonistic interaction, the assessment of the probable outcome of future encounters with the same individuals may be the best way of decreasing costs of fighting, but this may only be accomplished if animals are able to recognize individual conspecifics. We staged encounters between male lizards, Podarcis hispanica, to examine whether odoriferous cues are involved in rival recognition during agonistic interactions. We experimentally manipulated the odour of intruding males, creating familiar males with their own odour or bearing odours of unfamiliar males, and unfamiliar males bearing unfamiliar odours or odours of familiar males. The results showed that when familiar males were impregnated with scents from unfamiliar males, they elicited an aggressive response by resident unmanipulated males similar to that observed for a new unfamiliar male with unfamiliar odour. This suggests that resident males were unable to recognize familiar males when their own scents were removed. In contrast, responding males were less aggressive towards familiar males impregnated with their own odour and towards unfamiliar males impregnated with scents of familiar males, suggesting that when two males have already interacted, their scents become familiar for both males, and that the detection in successive encounters of the familiar scent suffices to reduce the aggressive response of territorial males. Therefore, recognition mechanisms based on chemical cues during agonistic encounters may contribute to reducing the intensity and the costs of fighting in P. hispanica and may play an important role in the organization of their social system.     

Voluntary corporate environmental initiatives and shareholder wealth

Researchers debate whether environmental investments reduce firm value or actually improve financial performance. We provide some compelling evidence on shareholder wealth effects of membership in voluntary environmental programs (VEPs). Companies announcing membership in EPA's Climate Leaders, a program targeting reductions in greenhouse gas emissions, experience significantly negative abnormal stock returns. The price decline is larger in firms with poor corporate governance structures, and for high market-to-book (i.e., high growth) firms. However, firms joining Ceres, a program involving more general environmental commitments, have insignificant announcement returns, as do portfolios of industry rivals. Overall, corporate commitments to reduce greenhouse gas emissions appear to conflict with firm value maximization. This has important implications for policies that rely on voluntary initiatives to address climate change. Further, we find that firms facing climate-related shareholder resolutions or firms with weak corporate governance standards – giving managers the discretion to make such voluntary environmentally responsible investment decisions – are more likely to join Climate Leaders; decisions that may result in lower firm value.     

Costs of switching social groups for dominant and subordinate dark-eyed juncos (Junco hyemalis)

I quantified the costs of switching from a familiar to an unfamiliar flock for captive dark-eyed juncos (Junco h. hyemalis) by measuring several physiological and behavioral variables before and after flock switching. Birds that were initially dominant dropped in status in unfamiliar flocks, and experienced increased metabolic rates, while subordinate birds appeared to undergo less physiological change when switching flocks. This difference occurred despite a lack of any rank-related differences in the effects of joining a new flock on rates of aggression, weight change, access to food, or plasma corticosterone levels. These results suggest that for dominant, but not subordinate, individuals there is a measurable metabolic cost to joining a new social group, even in the absence of adverse factors such as food limitation. Dominant individuals may be less likely than subordinates to leave familiar flocks because of their higher metabolic costs when joining a new social group.     

Voluntary corporate environmental initiatives and shareholder wealth

Researchers debate whether environmental investments reduce firm value or actually improve financial performance. We provide some compelling evidence on shareholder wealth effects of membership in voluntary environmental programs (VEPs). Companies announcing membership in EPA's Climate Leaders, a program targeting reductions in greenhouse gas emissions, experience significantly negative abnormal stock returns. The price decline is larger in firms with poor corporate governance structures, and for high market-to-book (i.e., high growth) firms. However, firms joining Ceres, a program involving more general environmental commitments, have insignificant announcement returns, as do portfolios of industry rivals. Overall, corporate commitments to reduce greenhouse gas emissions appear to conflict with firm value maximization. This has important implications for policies that rely on voluntary initiatives to address climate change. Further, we find that firms facing climate-related shareholder resolutions or firms with weak corporate governance standards – giving managers the discretion to make such voluntary environmentally responsible investment decisions – are more likely to join Climate Leaders; decisions that may result in lower firm value.     

Disentangling the effects of group size and density on shoaling decisions of three-spined sticklebacks (<Emphasis Type="Italic">Gasterosteus aculeatus</Emphasis>)

Many animals live in groups most of their life. One function of this behaviour is an increased predator protection whereas larger groups provide better protection than smaller ones. A causal explanation is that due to a higher number of shoal members the individual risk of being predated will decrease (“dilution effect”). Additionally, shoaling leads to increased predator confusion. This “confusion effect” can be strengthened by an increased group density, which often correlates with group size. Many studies found that individuals prefer the larger of two groups. However, whether this preference is due to a larger group size or because of an increased density of the larger group remained unclear. To disentangle these factors we gave three-spined sticklebacks (Gasterosteus aculeatus) the choice between shoals of (1) different group size and density, (2) different group size, but equal density and (3) equal group sizes, but different densities. As expected, test fish preferred the larger and denser shoal over the smaller, less dense one. This preference was lost when shoal size differed but density was kept constant. When shoal size was equal but density differed, test fish preferred the less dense shoal. However, this was only the case when test fish chose between two relatively dense shoals. On the other hand, when overall density was low, test fish did not discriminate between shoals of different densities. This result may be explained in terms of predator avoidance. The results show that shoaling preferences might not always be influenced by a higher number of group members but also by the density and cohesiveness of the respective groups. An erratum to this article can be found at     

Episodic targeting aggression and the histories of Lemur social groups

Summary Periodically, members of captive Lemur social groups target others for intense aggression. Over periods of several days, weeks, and sometimes months, one to three lemurs persistently follow and attack one or two particular group mates until the targets no longer associate with their group. Episodic targeting aggression is nonrandom with regard to time of year, group size and sex ratio, and the kinship, age, and gender of targets. The vast majority of episodes observed over the past 18 years at the Duke University Primate Center (DUPC) has occurred between like-sex adolescents and adults in conjunction with estrous cycling or infant births and most often after groups have reached apparent critical sizes. When unrelated adults have shared group membership, members of one family have almost invariably first targeted members of the other. In several groups, entire matrilines have gradually been evicted by members of another across periods of several years. When non-relatives have been absent, lemurs have evicted relatives that had previously formed separate subgroups. Episodic targeting aggression has been documented at the DUPC in over a dozen different social groups, comprising three different species. The phenomenon occurs repeatedly in groups held in a variety of large outdoor runs as well as in outdoor enclosures providing naturalistic space and physical structure. More-over, an appreciable number of recent observations in Madagascar suggest that the patterns we have documented well represent the phenomenon as it occurs in the wild. Targeting aggression based on group size, sex ratio, kinship and gender has been reported for no other primate taxon. We suggest that episodic targeting aggression reflects heretofore undescribed tactics in reproductive competition that may characterize many lemurid and indriid taxa. As such, the phenomenon has broad implications for the structure of lemur social groups and populations. Provisional models of the social dynamics and histories of Lemur social groups are presented for evaluation during upcoming field work.     

Habitat-specific chemical cues influence association preferences and shoal cohesion in fish

The structure of social animal groups can be dynamic, characterized by high rates of group fission and fusion. Despite this, group composition is often well ordered by factors such as species, body size and by numerous other phenotypic traits. Research in shoaling fishes has revealed that individuals refine group membership decisions still further and are capable of assimilating chemical cues pertaining to recent habitat and prey use by prospective group mates, preferring to associate with others whose recent resource use history closely matches their own. In this study, we firstly examined the dynamics of the formation and breakdown of these preferences, revealing that they can be acquired and replaced in a matter of just a few hours. Using such cues enables individuals to accurately assess the resource use of conspecifics, allowing them to indirectly sample the local environment while reducing the chances of acquiring outdated information that can precipitate maladaptive behaviors. Secondly, we found that shoals composed of individuals with shared recent habitat use history were more cohesive compared to those where the constituent individuals differed in recent habitat use. Increased shoal cohesion may reduce predation risk, and could enhance the ability of individuals to detect and use social information.     

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.     

Group size and foraging efficiency in yellow baboons

Summary I studied the foraging behaviour of adults in three different-sized groups of yellow baboons (Papio cynocephalus) at Amboseli National Park in Kenya to assess the relationship between group size and foraging efficiency in this species. Study groups ranged in size from 8 to 44 members; within each group, I collected feeding data for the dominant adult male, the highest ranking pregnant female, and the highest ranking female with a young infant. There were no significant differences between groups during the study in either the mean estimated energy value of the food ingested per day for each individual (385±27 kJ kg^-1 day^-1) or in the estimated energy expended to obtain that food (114±3 kJ kg^-1 day^-1). Mean foraging efficiency ratios, which reflect net energy gain per unit of foraging time, also did not vary as a function of the size of the group in which the baboons were living. There was substantial variation between days in the efficiency ratios of all animals; this was the result of large differences in energy intake rather than in the energy expended during foraging itself. The members of the smallest group spent on the average only one-half as much time feeding each day as did individuals in the two larger groups. However, they obtained almost as much energy while foraging, primarily because their rate of food intake while actually eating tended to be higher than the rate in the other groups. The baboons in the small group were observed closer to trees that they could climb to escape ground predators, and they also were more likely to sit in locations elevated above the ground while resting. Such differences would be expected if the members of the small group were less able to detect approaching predators than individuals that lived in the larger groups. The results of this study suggest that predator detection or avoidance, rather than increased foraging efficiency, may be the primary benefit of living in larger groups in this population.     

Unraveling fission-fusion dynamics: how subgroup properties and dyadic interactions influence individual decisions

Many species show fission-fusion group dynamics because it has clear advantages for flexibly exploiting heterogeneous environments. However, the mechanisms by which these dynamics arise are not well known. We used a hierarchical Bayesian model to disentangle the different influences on spider monkey (Ateles geoffroyi) individual fissions and fusions, including the three dimensions of fission-fusion dynamics (subgroup size, dispersion, and composition). Furthermore, we considered the influences of other individuals also leaving or joining a subgroup at the same time. We found that the most important influence on individual fissions and fusions is whether other individuals are also doing the same. Subgroup size and dispersion did not have clear effects on the probability that an individual fissioned or fusioned, while individuals tended to leave subgroups that were biased toward the opposite sex and to join subgroups that were biased toward their own sex. The networks constructed by the inter-individual influences during fissions and fusions were cohesive and did not show assortativity by sex or by degree. Individuals had a similar degree in both networks and each was influenced by a different set of individuals, suggesting a high fluidity in the social networks. We suggest that these networks reflect the way in which information about the environment flows as individuals follow one another during fissions and fusions.     

Individual dispersal status influences space use of conspecific residents in the common lizard, Lacerta vivipara

The effects of immigration on the behaviour of residents may have important implications for the local population characteristics. A manipulative laboratory experiment with yearlings of the common lizard (Lacerta vivipara) was performed to test whether the introduction of dispersing or philopatric individuals influences the short-term spacing behaviour of resident individuals. Staged encounters were carried out to induce interactions within dyads. The home cage of each responding individual was connected by a corridor to an unfamiliar “arrival cage” to measure the latency to leave their own home cage after each encounter. Our results showed that the time that pairs spent in close proximity was longer when a dispersing individual was introduced in the home cage. The latency to leave the home cage was longer after the introduction of a dispersing individual. These response variables were not influenced by the relative body sizes of contestants nor by the level of aggression towards each other. In contrast, the aggressive response was significantly influenced by the residency asymmetry established experimentally (“owner” of the home cage vs introduced individual). Our results suggest that the space use by resident individuals is influenced by the dispersal status of conspecifics. The potential ultimate causes driving this effect are discussed.     

Social discrimination tuning in ants: template formation and chemical similarity

To investigate the role of template plasticity in shaping nest-mate recognition processes in ants, we constructed experimental mixed-species groups of Manica rubida with either Myrmica rubra, Tetramorium bicarinatum or Formica selysi. Selecting Ma. rubida as the focal species, we observed the behaviour within mixed-species groups and the transfer rates of cuticular hydrocarbons (CHC) onto the focal ants, and we also tested the aggression of the focal species reared either alone or in association with each of the three different species. We show that Ma. rubida workers were always amicable towards their mixed group members, as towards members of the respective parental colonies, irrespective of the associated species. They did, however, express different levels of aggression towards single-species groups of the other species tested, depending on the species with which they were reared. The study suggests that similarity in CHC profiles in two species leads to a narrow template in mixed groups, while dissimilarity is followed by lower levels of aggression (a broader template), at least against species with similar CHC compound compositions (i.e. both a broader template in the focal ants and familiarity with the compound groups of the tested individuals operate together). This refutes the hypothesis that ants reared in mixed-species groups are systematically more tolerant. It also demonstrates that heterospecific information is not treated equally during development. We suggest that post-imaginal learning, template reforming and decision making are more precisely tuned when the two species' chemical complexes are similar.     

Intergroup conflict,ostracism, and the evolution of cooperation under free migration

Theoretical investigations and quasi-experimental evidence from modern conflict areas suggest that intergroup conflict and ostracism play a pivotal role in the evolution of cooperation. However, there is little direct evidence about the influence of intergroup conflict on human social behavior in the presence of endogenous group formation and unrestricted migration. This study introduces an experiment to examine the impact of intergroup conflict and ostracism on group formation, and human cooperation in a dynamic environment where group size, the occurrence of intergroup hostility and the threat of ostracism are endogenously determined. Here, we show that intergroup conflict may inhibit merging into single large human coalitions. The threat of ostracism is shown to increase the average group size and cooperation within the society. In addition, we find that competitive pressure between groups decreases the likelihood of social exclusion. Our results suggest that free migration between groups suppresses the impact of between-group competition on within-group cooperation. Moreover, our results stress the role of ostracism as a means to repress competition within groups and enhance group success in competition against other groups.     

Shark personalities? Repeatability of social network traits in a widely distributed predatory fish

Interest in animal personalities has generated a burgeoning literature on repeatability in individual traits such as boldness or exploration through time or across different contexts. Yet, repeatability can be influenced by the interactive social strategies of individuals, for example, consistent inter-individual variation in aggression is well documented. Previous work has largely focused on the social aspects of repeatability in animal behaviour by testing individuals in dyadic pairings. Under natural conditions, individuals interact in a heterogeneous polyadic network. However, the extent to which there is repeatability of social traits at this higher order network level remains unknown. Here, we provide the first empirical evidence of consistent and repeatable animal social networks. Using a model species of shark, a taxonomic group in which repeatability in behaviour has yet to be described, we repeatedly quantified the social networks of ten independent shark groups across different habitats, testing repeatability in individual network position under changing environments. To understand better the mechanisms behind repeatable social behaviour, we also explored the coupling between individual preferences for specific group sizes and social network position. We quantify repeatability in sharks by demonstrating that despite changes in aggregation measured at the group level, the social network position of individuals is consistent across treatments. Group size preferences were found to influence the social network position of individuals in small groups but less so for larger groups suggesting network structure, and thus, repeatability was driven by social preference over aggregation tendency.     

Patterns of use of space in a guild of territorial reef fishes

There exists on Heron Reef, at the southern end of the Great Barrier Reef, an 8-species guild of ecologically very similar, territorial, herbivorous reef fishes. All individuals of these species maintain territories on rubble substrata throughout juvenile and adult life. Territories are defended from all other guild members. Three rubble patches, each containing residents of 3 guild species: Pomacentrus apicalis, P. wardi, and Abudafduf lachrymatus, have been monitored for 12 to 18 months. This paper examines the patterns of use of space as shown by the antecedent and subsequent histories of sites chosen by 43 new colonists, and sites vacated through the loss of 34 residents on the rubble patches. In addition, variation through time is examined in the total amount of space held in territories on each rubble patch. The 3 species show similar preferences for space as colonists, although adult and juvenile colonists behave differently. Numbers of colonists detected on rubble patches are not proportional to the resident populations of the 3 species. Residents of the 3 species are equal in their abilities to enter spaces vacated through mortality, although they differ slightly in methods used to enter sites. They do not respond preferentially to sites previously occupied by any particular species. P. wardi shows a higher rate of mortality than the other species, and residents of this species are more often dislodged by new colonists. The total amount of space held on any rubble patch did not vary during the year from September, 1972 to October, 1973. The data are interpreted as supporting the hypothesis that the 3 species do not differ in their requirements for space on rubble patches, and that they are in competition for a short supply of such space. Differences exist in their strategies for obtaining and holding such space. These differences are important for explaining the continued presence of P. wardi on rubble patches. P. apicalis and a. lachrymatus are both specialists in holding territories on the upper reef slope. No differences have been detected in their requirements or competitive abilities. P. wardi is a fugitive species on the upper reef slope, coexisting because it maintains a refuge from competition by occupying some un-preferred sites off rubble patches.